Ornithological Monographs No. 45 Descriptions of Thirty-two New Species of Birds from the Hawaiian Islands: Part I. Non-Passeriformes Storrs L. Olson and Helen F. James Ornithological Monographs No. 46 Descriptions of Thirty-two New Species of Birds from the Hawaiian Islands: Part II. Passeriformes •y Helen F. James and Storrs L. Olson DESCRIPTIONS OF THIRTY-TWO NEW SPECIES OF BIRDS FROM THE HAWAIIAN ISLANDS: PART II. PASSERIFORMES ORNITHOLOGICAL MONOGRAPHS Edited by NED K. JOHNSON Museum of Vertebrate Zoology & Department of Integrafive Biology Life Sciences Building University of California Berkeley, California 94720 Ornithological Monographs, published by the American Ornithologists' Union, has been established for major papers too long for inclusion in the Union's journal, The Auk. Publication has been made possible through the generosity of the late Mrs. Carll Tucker and the Marcia Brady Tucker Foundation, Inc. Correspondence and manuscripts proposed for publication should be addressed to the Editor at the above address. Style and format should follow those of previous issues. Copies of Ornithological Monographs may be ordered from the Assistant to the Treasurer of the AOU, Max C. Thompson, Department of Biology, South- western College, 100 College St., Winfield, KS 67156. Price of Ornithological Monographs 45 and 46 bound together (not available separately): $25.00 prepaid ($22.50 to AOU members). Library of Congress Catalogue Card Number 91-72283 Primed by the Allen Press, Inc., Lawrence, Kansas 66044 Issued June 7, 1991 Copyright ¸ by the American Ornithologists' Union, 1991 ISBN: 0-935868-54-2 DESCRIPTIONS OF THIRTY-TWO NEW SPECIES OF BIRDS FROM THE HAWAIIAN ISLANDS: PART II. PASSERIFORMES. BY HELEN F. JAMES and STORRS L. OLSON Department of Vertebrate Zoology National Museum of Natural History Smithsonian Institution Washington, DC 20560 ORNITHOLOGICAL MONOGRAPHS NO. 46 PUBLISHED BY THE AMERICAN ORNITHOLOGISTS' UNION WASHINGTON, D.C. 1991 TABLE OF CONTENTS LIST OF FIGURES ............................................................................................................................................................ 4 LIST OF TABLES ................................................................................................................................................................. 5 INTRODUCTION ................................................................................................................................................................ 7 MATERIALS AND METHODS ......................................................................................................................... 8 SYSTEMATIC PALEONTOLOGY ................................................................................................................ 11 ORDER PASSERIFORMES .......................................................................................................................................... 11 Family Corvidae .............................................................................................................................................. 11 Genus Comus Linnaeus, 1758 ........................................................................ 11 Comus impluviatus, new species ................................................................ 12 Comus viriosus, new species .......................................................................... 19 Family Fringillidae ....................................................................................................................................... 22 Subfamily Carduelinae ............................................................................................................... 22 Tribe Drepanidini ...................................................................................................................... 22 Genus Telespiza Wilson, 1890 ....................................................................... 27 Telespiza persecutrix, new species .......................................................... 30 Telespiza ypsilon, new species ..................................................................... 35 Telespiza aft. ypsilon, Maui ............................................................................. 35 Genus Chloridops Wilson, 1888 ................................................................... 36 Chloridops sp., Kauai ............................................................................................... 36 Chloridops wahl new species ........................................................................ 36 Chloridops sp., Maui ................................................................................................ 40 Chloridops regiskongi, new species ........................................................ 40 Genus Incertae Sedis, Unassigned Maui Finch ........................... 43 Genus Rhodacanthis Rothschild, 1892 ............................................... 44 Rhodacanthis aft. palrneri Rothschild, 1892 ............................. 44 Rhodacanthis aft. fiaviceps Rothschild, 1892 ........................... 45 Orthiospiza, new genus ................................................................................................. 46 Orthiospiza howarthL new species ........................................................ 47 Xestospiza, new genus ................................................................................................... 52 Xestospiza conica, new species .................................................................... 53 Xestospizafastigialis, new species ......................................................... 55 Genus Incertae Sedis, Additional Oahu Finch ........................... 59 Genus Hemignathus Lichtenstein, 1839 .......................................... 59 Hemignathus upupirostris, new species ............................................ 60 Vangulifer, new genus .................................................................................................... 62 Vangulifer mirandus, new species ........................................................... 63 Vangulifer neophasis, new species .......................................................... 65 Aidemedia, new genus .................................................................................................... 66 Aidemedia chascax, new species ............................................................. 67 Aidemedia zanclops, new species ............................................................. 69 Aidemedia lutetiae, new species ................................................................ 71 Genus Ciridops Newton, 1892 ...................................................................... 73 Ciridops tenax, new species ............................................................................. 74 Ciridops sp., Oahu ....................................................................................................... 77 DISCUSSION ............................................................................................................................................................................... 78 ACKNOWLEDGMENTS ............................................................................................................................................. 83 SUMMARY ..................................................................................................................................................................................... 85 LITERATURE CITED ..................................................................................................................................................... 86 Figure 1. 2. 3. 4. 5. 6. 7. 8. 9. LIST OF FIGURES Diagram showing measurements of the maxilla ................................. 9 Diagram showing measurements of the mandible ............................... 10 Skulls and mandibles of Hawaiian Corvus in lateral view ........ 13 Skulls of Hawaiian Corvus in dorsal view ....................................................... 14 Skulls of Hawaiian Corvus in ventral view .................................................... 15 Mandibles of Hawaiian Corvus in dorsal view ......................................... 17 Humeri of Hawaiian Corvus .............................................................................................. 20 Hindlimb elements of Hawaiian Corvus ........................................................... 21 The ventral maxillae in finch-billed drepanidines and Loxops virens ................................................................................................................................................................. 26 10. Box plots of measurements of the maxilla in Telespiza ............... 31 11. Maxillae of Telespiza .................................................................................................................. 32 12. Mandibles of Telespiza in lateral view ................................................................ 33 13. Mandibles of Telespiza in dorsal view ................................................................ 34 14. Maxillae of Chloridops ............................................................................................................. 37 15. Mandibles of Chloridops in dorsal view ............................................................ 38 16. Maxillae and mandibles of Chloridops wahi and C. kona in lateral view ................................................................................................................................................ 42 17. Maxillae and mandible of Chloridops regiskongi in lateral view 43 18. Cranium, palatine, and quadrate of Orthiospiza howarthi 47 19. Maxillae of Orthiospiza howarthi and Rhodacanthis fiaviceps 48 20. Mandibles of Orthiospiza howarthi and Rhodacanthis fiaviceps 50 21. Postcranial elements of Orthiospiza howarthi ............................................ 51 22. Maxillae of Melamprosops and Xestospiza .................................................... 54 23. Mandibles of Melamprosops and Xestospiza ............................................... 56 24. Bones of the holotype of Xestospizafastigialis ......................................... 58 25. Mandibles of Hemignathus ................................................................................................. 61 26. Maxillae of Vangulifer ............................................................................................................... 64 27. Mandibles of Vangulifer .......................................................................................................... 65 28. Maxillae of Aidemedia and Loxops sagittirostris .................................. 68 29. Mandibles of Aidemedia and Loxops sagitt•rostris in lateral view .................................................................................................................................................................... 70 30. Mandibles of Aidemedia and Loxops sagittirostris in dorsal view ..................................................................................................................................................................... 72 31. Maxillae of Ciridops in lateral view ......................................................................... 74 32. Maxillae of Ciridops in dorsal and ventral views .................................. 75 33. Mandibles of Ciridops in dorsal view ................................................................... 77 34. Mandibles of Ciridops in lateral view ................................................................... 78 35. Pelvic and hindlimb elements of Ciridops ...................................................... 79 LIST OF TABLES Table 1. 2. 3. 4. 5. 6. 7. 8. 9. 10. 11. 12. 13. 14. Measurements of major skeletal elements of Hawaiian Corvus. 18 Measurements of the maxilla in Telespiza and Loxioides ................. 28 Measurements of the mandible in Telespiza and Loxioides ........... 29 Distribution of species of Telespiza .............................................................................. 30 Measurements of the maxilla in Chloridops ......................................................... 39 Measurements of the mandible in Chloridops ................................................... 39 Measurements of the maxilla in Rhodacanthis and Orthiospiza 45 Measurements of the mandible in Rhodacanthis and Orthiospiza 45 Measurements of the maxilla in Xestospiza, Melarnprosops and Psittirostra .......................................................................................................................................................... 53 Measurements of the mandible in Xestospiza, Melarnprosops and Psittirostra ......................................................................................................................................................... 57 Measurements of the maxilla and mandible in Aidemedia and Loxops sagittirostris .............................................................................................................................. 69 Measurements of the maxilla and mandible in Ciridops .................... 75 Measurements of the postcranial skeleton in Ciridops ........................... 76 Fossil and historical distribution of endemic passerine birds in the Hawaiian Archipelago ............................................................................................................. 80 INTRODUCTION The present work continues the description of new species of birds from the Hawaiian Islands that have come to light through fossil collecting over the past two decades (Olson and James 1982a, b; 1984). In the preceding part (Olson and James, 1991), we briefly describe the various fossil localities and introduce 3 new genera and 16 new species ofnon-passerines, including 1 petrel, 5 raptorial species, and 10 species that were flightless or nearly so. Here we treat the Passeriformes, describing as new 2 species of crows (Corvidae) and 4 genera and 14 species of Hawaiian finches (Drepanidini). We also discuss but do not name an additional 8 possible new species that are as yet known from insufficient material for proper diagnosis. Although we have found new fossil species of Meliphagidae as well, we have postponed describing them pending further revisionary work. Also not treated at this time are fossils of the families Myiagridae and Muscicapidae, which do not appear to contain any new taxa although they have not been thoroughly studied. The new passerine taxa are derived from rich fossil deposits found on the islands of Kauai, Oahu, Molokai, and Maui. The less numerous passerine fossils found mainly in archaeological contexts on the island of Hawaii do not include any new taxa, and no passerine fossils have yet been found on the other main islands (Niihau, Lanai, Kahoolawe). Small passerine bones usually have a different taphonomic history from those of crows and the larger non-passerines that often occur in the same deposits. Passerines from aeolian dunes on Kauai and Molokai were deposited as pellets cast by extinct owls of the genus Grallistrix, which evidently roosted either in dune shrubs or directly on the ground in hollows in the dunes. The only non- passerine that is abundant in such owl pellet deposits is the tiny flightless rail Porzana menehune of Molokai. Bones of crows and larger non-passerines in the same deposits are thought to be remains of individuals that died among the dunes and were buried by shifting sands, or were deposited either in burrows of nesting seabirds or as human midden material. Near Barbers Point on Oahu, fossils were found in sediments that had accu- mulated in sinkholes in a raised limestone reef. In many of the smaller sinkholes, passerines are rare and may have resulted from the chance death of an individual in or over the sink. In other sites, passerine bones are more abundant and may have originated in pellets from nearby roosts of Grallistrix. Even in these presumed owl pellet deposits, passerine remains are less concentrated than is usual in primary owl roost accumulations such as those in the dunes mentioned previously, or in barn owl (Tyro spp.) roosts in West Indian caves (Pregill 1981). It may be that the owl pellet material at Barbers Point was transported a short distance and redeposited by water, resulting in less concentrated deposits. On Maul, fossils were found in lava tubes, many of which are concentrated along the southwest rift zone of Mr. Haleakala. The most important of these sites is Puu Naio Cave (James et al. 1987), where finely stratified Holocene sediments were partially excavated in 1984 and 1988. Passerine remains here are believed to have been concentrated in the pitfall opening of the cave by individuals of Grallistrix roosting on ledges. During floods, some of the prey remains were transported into the cave where they were preserved. 8 ORNITHOLOGICAL MONOGRAPHS NO. 46 Passefines must occasionally penetrate caves beyond the light zone, probably at dusk, become disoriented, and never find their way back out, which leads to their associated skeletons being left in a circumscribed area on the floor of the cave. Outstanding examples of this mode of deposition are the flooded cavern at Barbers Point (Site 50-Oa-B6-139), where exquisitely preserved associated skel- etons of crows and a species of meliphagid were recovered with SCUBA gear, and Crystal Cave on Maul, where the situation of the cave mouth in the wall of a steep gulch makes it accessible only to bats and volant birds, several species of which were found inside. The following descriptions do not include a heading for the geological age of the fossils, which is Holocene for all of the new drepanidines, and Late Quaternary for the new crows. A less precise age is given for the crows because both new species occur in the flooded cavern at Barbers Point, mentioned above. Fossils from this site, thought to be older than those from sediment-filled sinkholes in the vicinity, possibly date to the Late Pleistocene. The limited amount of radiocarbon data that is available from the dune sites on Kauai and Molokai, and from the sediment-filled sinkholes on Oahu, indicates that all of the passerine fossils from these sites are probably less than 7,000 years old, and most are less than 5,000 years old (Olson and James 1982b; Olson and James 1991). The bones that have been dated from lava tube sites on Maul are all less than 5,500 years old (Thomas W. Stafford, Jr., pers. comm.), except that those from Puu Naio Cave span the Holocene from at least 8,000 years ago to the present (James et al. 1987), and those from the Crystal Cave site, which have not been dated, may be even older. The only passefine fossils that are certainly known to antedate the Holocene are those from Ulupau Head, Oahu, where the bone-beating lacustrine and col- luvial sediments are more than 120,000 years old (James 1987). This site is still being actively collected and the material has not been fully studied. MATERIALS AND METHODS Format and organization are as in Part I (Olson and James 1991). Within genera, or groups of related species within a genus, the order of presentation in text and tables is geographic, from west to east. For economy of space, not all paratypes are listed (see Olson and James 1991). Although statements of distri- bution include species identified from the Pleistocene deposits at Ulupau Head, none of the specimens from that locality were used in the diagnoses of new taxa or are considered paratypes. One of the greatest obstacles to carrying out this study was the lack of adequate comparative osteological specimens of the endemic species that make up the historically known passefine fauna. Many of these species became extinct or endangered before any skeletal specimens of them were ever prepared. We were able to compensate for this by removing the skull and mandible, and sometimes limb elements, from study skins by the method described by Olson et al. (1987), so that at least one skull and mandible were available for all but two of the species of drepanidines. Measurements were taken with digital calipers to the nearest 0.1 mm. Because of the highly diagnostic morphology of the maxilla and mandible, 23 different measurements were taken of these elements, as shown in Figs. 1 and 2. These NEW HAWAIIAN BIRDS 9 3 11 8 2 5 10 ' 4 Flo. 1. Diagram showing measurements of the maxilla (Loxioides bailleui in dorsal, ventral, and lateral views [top to bottom]): 1, dorsal length; 2, ventral length; 3, length from lateral comer of nasofrontal hinge; 4, length fromjugal articulation; 5, length from anterior narial opening; 6, maximum width; 7, width of nasofrontal hinge; 8, length of narial opening; 9, height of narial opening; 10, height through lateral nasal bar; 11, minimum width of dorsal nasal bar. were supplemented with measurements of the cranium and postcranial skeleton when such elements were preserved and could be positively identified. Mensural data are organized to serve the dual purpose of characterizing the hypodigm of each new species and of identifying the holotype itself by its unique dimensions. Tables give means, standard deviations, and ranges for sample sizes of 5 or greater, with the measurement of the holotype listed separately in the text, but list indi- 10 ORNITHOLOGICAL MONOGRAPHS NO. 46 9 8 4 7 11 i 12 F•a. 2. Diagram showing measurements ofthe mandible (Vestiaria coccinea, dorsal view, above; Loxioides bailleui, dorsal and lateral views, middle and below): 1, total length; 2, length of tomial crest; 3, symphysis length; 4, greatest width of symphysis; 5,greatest width of mandible; 6, symphysis height; 7, length of lateral cotyla; 8, length of lateral cotyla plus retroarticular process; 9, width of articular end with medial process; 10, height at lateral cotyla; 11; height at angle of mandible; 12, ramus length (middle part). NEW HAWAIIAN BIRDS 11 vidual data for samples of 4 or fewer. In the latter case, the measurement of the holotype is identified in the table with an H. SYSTAT (Wilkinson 1989) and SYGRAPH (Wilkinson 1988) software were used for statistical tests and box plots. Terminology is drawn from Baumel et al. (1979), Richards and Bock (1973), Howard (1929), and Zusi (1978; the term cranial fenestra only). We employ the terms of position and direction used by Howard (1929), instead of those rec- ommended by Baumel et al. (1979). Thus, in place of cranial, rostral, and anterior, we use anterior, and in place of caudal we use posterior. As used here, maxilla and mandible refer to the bony elements of the jaws, whereas maxillary rostrum and mandibular rostrum refer to the external rham- phothecal covering together with the underlying bony parts of these elements. Narial opening refers specifically to the external margin of the bony naris. The mandibular ramus between the articular end and the symphysis is referred to as the middle part of the ramus, anglicized from Baumel et al. (1979) (see Fig. 2). All of the Hawaiian fossil material treated in this study is housed either in the Department of Zoology, Bernice P. Bishop Museum, Honolulu, or the Department of Palcobiology, National Museum of Natural History, Smithsonian Institution, Washington, D.C. Comparative material was examined from the following col- lections: AMNH--American Museum of Natural History, New York. ANSP--Academy of Natural Sciences, Philadelphia. BMNH--British Museum (Natural History), Tring. BBM, BBM-X, BPBM--Bernice P. Bishop Museum, Honolulu. AU--Geology Department, Auckland University, Auckland. CMC--Canterbury Museum, Christchurch. MCZ--Museum of Comparative Zoology, Harvard University, Cambridge, Mas- sachusetts. MVZ--Museum of Vertebrate Zoology, University of California, Berkeley. NMNZ--National Museum of New Zealand, Wellington. PB--Pierce Brodkorb Collection, University of Florida, Gainesville. USNM--National Museum of Natural History, Smithsonian Institution, Wash- ington, D.C. ZMB--Museum f'dr Naturkunde, Humboldt-Universi•t, Berlin. SYSTEMATIC PALEONTOLOGY Order PASSERIFORMES Family CORVIDAE Genus Corvus Linnaeus, 1758 Two new species of crows in lowland fossil sites on Oahu and Molokai (Olson and James 1982a, b, 1984) are known from associated fossil skeletons preserved so well as to rival the best skeletal preparations in modern museum collections. These fossils were collected under water in a flooded cavern near Barbers Point, Oahu. We have designated specimens from this cavern as the holotypes for both of the new species, although one of the species is known so far from Oahu only from the holotype, and occurs far more abundantly in the dune sites on Molokai. Bones of crows have also been found in lava tubes on Maui (James et al. 1987) 12 ORNITHOLOGICAL MONOGRAPHS NO. 46 and Hawaii. These are all isolated elements or fragmentary skeletons that need further study before they can be identified to species with reasonable assurance. We compared the new Hawaiian crows to all other species of Corvus that occur in the Pacific Basin and peripheral continental areas, except two geographically restricted species for which specimens were not available. To simplify our diagnoses, we can eliminate the following species from further discussion. The new Hawaiian crows are markedly larger than Corvus dauuricus, C. moneduloides, C. enca, C. typicus, C. kubaryi (bill also narrower), C. frugilegus, C. brachyrhynchos, C. caurinus, C. imparatus, C. orru, C. torquatus (bill also less deep), or C. cryptoleucus. The bills of the new species are deeper than in C. coronoides, C. mellori, C. torquatus, or C. validus. Compared to the new species, Corvus corone has the ventral surface of the maxilla less excavated, the nostril longer, and the bill less deep; C. bennetti has a smaller bill; and C. tristis has the mandible less deep and the sternum relatively large in proportion to humerus length. Comparative material examined: The complete skeleton of Corvus hawaiiensis that we used in our comparisons, USNM 501638, is from an atypically small male bird. We supplemented this specimen by vemoving the skull, mandible, and several long bones from the skin of a large male ofC. hawaiiensis, USNM 177993. Specimens of C. fiorensis (restricted to Flores in the Lesser Sundas) and C. fusci- capillus (restricted to parts of New Guinea and its satellites) were not available for comparison. Skeletal material from the Smithsonian collections included: C. dauuricus male, 292083; females, 292082, 319401. C. moneduloides male, 561635; female, 561634. C. enca male, 225830; unsexed, 224802. C. typicus male, 226205 (trunk only). C. kubaryi unsexed, 613280. C. validus males, 557299, 557300, 558297; females, 489028, 557301. C. tristis female, 489028. C. frugilegus males, 290314, 291673. C. brachyrhynchos male, 554206; female, 499510. C. caurinus males, 561899, 612996; females, 612993, 612995. C. corone male, 500773; un- sexed, 289948. C. macrorhynchos males, 290456, 292081, 500768; females, 290955, 500774; unsexed, 318366. C. orru male, 559044, females, 558338, 559045. C. torquatus males, 289947, 291412, 292078; female, 292858. C. cryptoleucus males, 553971, 553972, 553973, 555254; females, 498679, 554139. C. corax males, 489704, 499938; female, 555261; unsexed, 18622, 290441. Of C. mo- riorum, we examined the following subfossil specimens: NMNZ S 962, skull and partial skeleton; AU 6120, partial postcranial skeleton; AU 6121.16, mandible; CMC AV3310, skull and partial skeleton. We also made comparisons with study skins of the following species in the Smithsonian collections: C. hawaiiensis (12 specimens), C. woodfordi, C. meeki, C. imparatus, C. bennettL C. coronoides, and C. mellori. Corvus impluvlatus, new species (Figs. 3B, 4B, 5B, 6B, 7B, 8B, E) "Corvus, deep-billed" Olson and James, 1982b:38, 44; 1984:771. Holotype: Nearly complete skeleton, BBM-X 153652. Collected 22 July 1977 by Storrs L. Olson, Helen F. James, Aki Sinoto and others, with the aid of SCUBA gear. The specimen includes the cranium with the maxilla, palatines, and jugals in place (Figs. 3B, 4B, 5B), both pteryg0ids, both quadrates, the left prefrontal, NEW HAWAIIAN BIRDS 13 B FiG. 3. Skulls and mandibles of Hawaiian Corvus in lateral view. A, C. hawaiiensis (skull, USNM 177993; mandible, USNM 501638); B, C. impluviatus, new species, holotype (BBM-X 153652); C, C. viriosus, new species, holotype (USNM 386435; image of cranium is of the left side printed in reverse). Scale = 3 cm. 14 ORNITHOLOGICAL MONOGRAPHS NO 46 B F•G. 4. Skulls of Hawaiian Corvus in dorsal view. ,4, C. hawaiiensis (USNM 177993); B, C. •mpluviatus, new species, holotype (BBM-X 153652); C, C. v•nosus, new species, holotype (USNM 386435). Scale = 3 cm. NEW HAWAIIAN BIRDS 15 B FIG. 5. Skulls of Hawaiian Corvus in ventral view. A, C. hawaiiensis (USNM 501638); B, C. impluviatus, new species, holotype (BBM-X 153652); C, C. viriosus, new species, holotype (USNM 386435). Scale = 3 cm. 16 ORNITHOLOGICAL MONOGRAPHS NO. 46 the mandible (Figs. 3B, 6B), twelve tracheal rings, the syringeal drum, the furcula, the sternum, ten complete ribs plus some fragments, ten cervical vertebrae (the complete cervical series except for the fourth vertebra), three caudal vertebrae, all of the long bones of the pectoral skeleton (scapulae, coracoids, humeri (Fig. 7B), ulnae, radii, carpometacarpi), the radiale, ulnare, and one alar phalanx from the right side, the left femur (Fig. 8B), both tibiotarsi and fibulae, the right tar- sometatarsus (Fig. 8E), and twelve pedal phalanges. Type locality: Flooded cavern, Site 50-Oa-B6-139, Barbers Point, Oahu, Ha- waiian Islands. Distribution: Oahu: Barbers Point. Etymology: An adjective formed from Latin, impluvium, n., the skylight in the roof of the atrium of a Roman house, in reference to the nature of the type locality, which we entered by leaping through a skylight into the clear water below. The root word pluvius, meaning rainy, may be taken as a further allusion to the watery environment of the cavern. Measurements (ram) ofholotype: Maxilla: dorsal length, 66.8; length from an- terior rim of narial opening, 40.2; maximum width, 26.7; width of nasofrontal hinge, 23.8; maximum height, 21.4; height through lateral nasal bar, 24.1; length of narial opening, 13.7; height of narial opening, 8.5. Cranium: total length with maxilla, 105.1; length from supraoccipital to nasofrontal hinge, 45.4; length from nasofrontal hinge to bill tip, 66.7; orbit length (ectethmoid to postorbital process), 26.2; width of frontal between orbit rims, 25.6; width behind postorbital pro- cesses, 42.9; height from basitemporal plate to sloall roof, 31.7; length ofbasitem- poral plate (anterior margin of basitemporal plate to posterior extremity of oc- cipital condyle), 15.7; width between external rims of the articular faces for the quadrates, 39.9. Mandible: total length, 87.3; length of tomial crest, 52.8; sym- physis length, 22.2; greatest width of mandible, 45.0; width of articular end with medial process, 16.9; height at angle of mandible, 15.5; greatest height of sub- rhamphothecal ramus, 13.8. Scapula: length, 57.2; proximal width, 14.2. Sternum: length through manubrial spine, 62.1; width below costal facets, 32.2; length of carina, 60.5; depth of carina, 18.6. Coracoid: length, 51.2; width of sternal end, 13.2. Humerus: length, 78.5; proximal width, 21.2; length of deltoid crest, 24.3; mid-shaft width, 6.9; distal width, 17.1. Ulna: proximal width, 11.8; distal width, 11.2. Radius: length, 85.9. Carpometacarpus: length, 55.6; proximal depth, 12.9; distal depth, 12.0. Pelvis: xvidth across trochanters, 40.5. Femur: length, 63.1; proximal width, 13.7; mid-shaft depth, 4.6; distal width, 13.2. Tibiotarsus: length without cnemial crest, 105.4; proximal width, 32.9; length of fibular crest, 17.8; distal width, 11.4. Tarsometatarsus: length, 72.2; proximal width, 12.7; mid-shaft width, 4.6; depth of third trochlea, 5.2. For additional measurements, see Table 1. Paratypes: Partial skeleton, USNM 386431, including the cranium (with max- illa, palatines, and jugals attached), the right pterygoid, the left quadrate, the mandible, the furcula, sternum and pelvis, fifteen complete ribs plus some rib fragments, the atlas and axis and seven additional cervical vertebrae, four thoracic vertebrae, the right scapula, the left coracoid, the humeri, radii, carpometacarpi, and the first phalanges from the right and left major digit, all of the long bones of the hindlimb (femora, tibiotarsi, fibulae, tarsometatarsi), and the first phalanx of the hallux. NEW HAWAIIAN BIRDS 1 ? B FIG. 6. Mandibles of Hawaiian Corvus in dorsal view. A, C. hawaiiensis (USNM 501638); B, C. irnpluviatus new species, holotype (BBM-X 153652); C. C. viriosus new species, holotype (USNM 386435). Scale = 3 cm. Partial skeleton, USNM 386428, including the skull and mandible, right cor- acoid, eroded sternum and pelvis, some ribs and vertebrae, and the paired scap- ulae, humeri, ulnae, radii, femora, tibiotarsi, and tarsometatarsi. Partial skeleton, USNM 386427, including the mandible, furcula, sternum, 18 ORNITHOLOGICAL MONOGRAPHS NO. 46 TABLE 1 L•x•o•-• MEASUREMEN'mS (MM) or THE MAJOR SKELETAL ELEMENTS IN HAWAIIAN Corvus. MEAN, STANDARD DEVIATION, AND RANGE ARE GIVEN FOR SAMPLE S•z• GRF_ATER THAN 4; DATA ARE LISTED INDMDUALLY FOR SAMPLE SIZES OF 4 OR FEWER. H = HOLOTYPE Corpus Corvus impluviatus Corvu* viriosus hawaiiensis Skull 102.3 -+ 4.6 95.1-107.4 n=6 Quadrate 16.1, 16.1, 16.2 H Maxilla 64.7 -+ 3.4 59.6-68.5 n=6 Mandible 86.5 -+ 3.1 80.7-89.4 n=8 Sternum 62.1 _+ 2.5 59.3-66.1 n=5 Synsacrum 52.2 H, 55.8, 57.9, 58.3 Scapula 56.9, 57.2 H, 60.2 Coracoid 50.8 ñ 1.8 47.9-52.4 n=5 105.9, 112.6 H 84.3, 94.0 15.0, 15.0, 15.5 11.4, 13.3 65.1, 74.2 H 47.2, 57.2 83.0, 89.8, 94.5 H 67.8, 77.1 57.8 H 49.8, 51.0 H 45.9 57.0 47.5 45.1, 45.2, 46.2, 49.2 H 40.5 Humerus 79.1 ñ 1.8 73.4 + 2.9 62.9 76.3-80.7 70.8-77.7 n=5 n=8 Ulna 90.8, 94.2 H, 94.9, 96.7 85.3 _+ 3.6 72.5, 80.6 77.4-88.9 n=11 Radius 85.9 ñ 2.3 78.5 H, 80.3, 80.3, 81.0 66.6 82.2-87.8 Carpometacarpus 55.4 ñ 1.4 50.8, 50.9 H, 52.8 43.5 53.3-57.2 n=5 Femur 62.8 ñ 2.3 60.4 _+ 2.2 52.3 59.6-66.1 58.2-63.4 n=5 n=8 Tibiotarsus without 104.1 ñ 3.6 98.7, 99.2, 102.2 H, 106.2 88.4, 97.9 cnemial crest 98.5-108.3 n=5 Tarsometatarsus 71.4 ñ 2.9 69.1 _+ 3.3 61.4, 69.0 67.4-75.1 66.2-76.2 n=5 n=7 pelvis, a vertebra and a fib, and the long bones of the pectoral skeleton from the left side. Partial skeleton, BBM-X 153649, including the skull, palatines, pterygoids, quadrates, mandible, furcula, sternum, pelvis, some fibs, vertebrae, and tracheal tings, all of the long bones of the pectoral skeleton except one radius, all of the long bones of the hindlimb except one fibula, plus two pedal and one ungual phalanges. NEW HAWAIIAN BIRDS 19 Partial skeleton, BBM-X 153654, including the skull, mandible, eroded left humerus, and the femora and tarsometatarsi. Partial skeleton, BBM-X 153648, including the skull and mandible, furcula, eroded sternum and pelvis, and some bone fragments. Associated mandible and tibiotarsus, BBM-X 153651. Measurements of Paratypes: See Table 1. Diagnosis: A large species of Corvus with a high, arched bill. The bill is broader and deeper than in C. woodfordi or C. meeki. Compared to C. macrorhynchus, C. corax, and C. moriorum, the nostril is less elongated anteroposteriorly, the membranes in the nasal cavities are more extensively ossified, and the interorbital septurn is not fenestrated. C. impluviatus differs further from C. macrorhynchus in having the bill shorter and deeper; from C. corax in having the bill deeper and the dorsal nasal bar broader; and from C. moriorum in having the bill more arched, the cranial fenestra smaller or absent, and the articular end of the mandible larger. Compared to C. hawaiiensis and C. viriosus, new species, the maxilla is deeper and more arched, the dorsal and lateral nasal bars are wider, the ossification of membranes in the nasal cavities is more extensive (including ossification of the nasal septum, floor of the nasal cavity, and the partial occlusion of the narial openings), the zygomatic process is broader, the olecranon is slimmer, and the posterior projection of the ilium is longer and slimmer. C. impluviatus differs further from C. viriosus but agrees with C. hawaiiensis in having a relatively short bill, with a less excavated ventral surface of the maxilla, a broader frontal, broad postorbital processes, broad and rounded rather than square-tipped transpalatine processes, and the mandible with a shorter symphysis, a more decurved tomial crest, and with a slight ventral projection to the posterior fossa that is not de- veloped in C. viriosus. Remarks: The postcranial bones of C. impluviatus average longer than those of C. hawaiiensis or C. viriosus, although the average size difference between C. impluviatus and C. viriosus is generally slight (Table 1). The short, arched bill and the extensive ossification within the nasal and orbital cavities are very distinctive characters of this spedes. See further remarks under the following species. Corvus viriosus, new species (Figs. 3C, 4C, 5C, 6C, 7C, 8C, F) "Corvus sp., slender-billed" Olson and James, 1982b:38, 44; 1984:771. Holotype: Partial skeleton, USNM 386435. Collected 26 July 1977 by Storrs L. Olson. The specimen includes the cranium with the maxilla attached (Figs. 3C, 4C, 5C), the palatines, the mandible (Figs. 3C, 6C), the furcula, sternum, and pelvis, four complete ribs and some rib fragments, two thoracic and one caudal vertebrae plus the pygostyle, the right humerus (Fig. 7C), ulna, and carpometa- carpus; the left coracoid, radius and first phalanx of the major alar digit; the left femur (Fig. 8C), and both tibiotarsi and tarsometatarsi (Fig. 8F). The bones, which are darker in color and more striated than most other crow bones from the site, are in nearly perfect condition. The skull of this bird was found on the floor of the cavern near the skull and other bones of a specimen of C. impluviatus..The mandible and postcranial bones were retrieved from a nearby hole and crevice into which they had slipped. 20 ORNITHOLOGICAL MONOGRAPHS NO 46 A B C FIG. 7. Humeri of Hawaiian Corvus in anconal view. A, C. hawaitensis (USNM 501638); B, C. irnpluviatus, new species, holotype (BBM-X 153652); C, C. viriosus, new species, holot•pe (USNM 386435). Scale = 3 cm. Type locality: Flooded cavern, Site 50-Oa-B6-139, Barbers Point, Oahu, Ha- waiian Islands. Distribution: Oahu: Barbers Point (holotype only). Molokai: Moomomi dunes and Ilio Point. Etymology: Latin, viriosus, robust, strong; from the larger size of this species compared to C. hawaiiensis and from the sturdy construction of the cranium and mandible of Hawaiian corvids generally. Measurements (ram) ofholotype: Maxilla: length from anterior narial opening, 46.5; maximum width, 27.0; width of nasofrontal hinge, 22.8; height through lateral nasal bar, 23.3; length of narial opening, 17.6; height of narial opening, 9.6. Cranium: length from supraoccipital to nasofrontal hinge, 44.0; length from supraoccipital to bill tip, i12.6; orbit length (ectethmoid to postorbital process), 24.3; width of frontal between orbit rims, 21.3; width posterior to postorbital processes, 41.0; height from basitemporal plate to skull roof, 28.7; length of basitemporal plate {anterior margin of basitemporal plate to posterior extremity of occipital condyle), 14.7; width between external rims of the articular faces for the quadrates, 38.2. Mandible: length of tomial crest, 56.4; symphysis length, 28.2; greatest width of mandible, 42.3; width ofarticular end with medial process, 16.2; height at angle of mandible, 14.0; greatest height of sub-rhamphothecal ramus, 13.2. NEW HAWAIIAN BIRDS 21 A B C D E F F•G. 8. Bones of the hindlimb of Hawaiian Comus. A, B, C, left femora in posterior view; D, E, F, right tarsometatarsi n anterior view; A. C. hawaiiensis, small male (USNM 501638); D, C. ha- waiiensis, large male (USNM 177993); B, E, C. irnœ1uviatus, new species, holotype (BBM-X 153652); C, F, C. viriosus, new species, holotype (USNM 386435). Scale = 3 cm. Sternum: width below costal facets, 30.3; length of carina, 56.8; depth of carina, 17.4. Coracoid: width of sternal end, 13.2. Humerus: length, 76.1; proximal width, 20.0; length ofdeltoid crest, 18.6; mid-shaft width, 6.4; distal width, 18.0. Ulna: length, 86.8; proximal width, 11.6; distal width, 10.4. Carpometacarpus: proximal depth, 12.8; distal depth, 11.8. Major alar digit, phalanx 1: length, 22.5. Pelvis: width across antitrochanters, 38.1. Femur: length, 61.9; proximal width, 12.9; mid-shaft depth, 5.1; distal width, 13.9. Tibiotarsus: proximai width, 17.3; length from proximal articulation to distal fibular crest, 31.4; distal width, 11.4. Tar- sometatarsus: length, 68.0; proximal width, 11.9; mid-shaft width, 4.8; depth of third trochlea, 5.4. Paratype: Molokai: damaged cranium and associated bones, BBM-X 148156, including part of the fused parietals and frontal with the maxilla attached, the prepalatine bars, the quadrate, and the mandible lacking the right articular end and half of the right dentary. Measurements ofparatype.' Included in Table 1, along with measurements of other specimens not listed as paratypes. Diagnosis: A large species of Corvus with a long, straight bill. The bill is straighter and the dorsal nasal bar is narrower than in C. woodfordi. The mandibular ramus is deeper and the tarsometatarsus is shorter than in C. meeki. The interorbital 22 ORNITHOLOGICAL MONOGRAPHS NO. 46 fenestra is smaller and the narial opening is less elongate anteroposteriorly than in C. macrorhynchus, C. corax, or C. moriorum. C. viriosus differs further from C. macrorhynchus in having the maxillary rostrum less arched anteriorly; from C. corax in having the bill deeper, the dorsal nasal bar broader, and the membranes in the nasal cavities more extensively ossified; and from C. moriorum in having the cranial fenestra smaller, the ossification of membranes in the nasal cavities more extensive, the mandibular ramus deeper, and the articular end of the man- dible larger. This species differs from C. hawaiiensis and C. impluviatus in having the bill longer, straighter, and less deep (intermediate in C. hawaiiensis), with more pro- nounced excavation of the ventral maxilla, narrower dorsal and lateral nasal bars (also intermediate in C. hawaiiensis), a longer mandibular symphysis, the tomial crest of the mandible nearly straight (gradually curved in C. hawaiiensis and C. impluviatus), the slight ventral projection of the posterior fossa of the mandible absent, the frontal less broad, the postorbital process slimmer, and the transpala- tine process square-tipped (broad and rounded in C. hawaiiensis and C. implu- viatus). The olecranal fossa of the humerus lacks the deep, rounded pit that is present in C. hawaiiensis and C. impluviatus. C. viriosus differs further from C. impluviatus, but agrees with C. hawaiiensis, in having a narrower dorsal nasal bar, a relatively slim zygomatic process, a stouter olecranon, and a shorter and broader posterior projection of the ilium. Remarks: The most obvious way to distinguish the three Hawaiian species of Corvus is by bill shape: C. hawaiiensis has a short, moderately arched bill; C. impluviatus has a high, impressively arched bill; and C. viriosus has a long, rel- atively straight bill. On average, the long bones of C. viriosus are intermediate in length between the smaller C. hawaiiensis and the larger C. impluviatus, although there is some overlap with either species (Table 1). The osteological comparisons we made for the purpose ofdiagnosing new spe- cies were not sufficiently comprehensive to determine the phylogenetic position of the Hawaiian crows relative to species of Corvus outside the archipelago. Our impression is that the Hawaiian crows probably did not arise from Nearctic ancestors but may very well have been derived from Australasia, much like Chasiernpis and the Hawaiian Meliphagidae. We are less confident than previously (Olson and James 1982b:51) that more than one colonizing species gave rise to the Hawaiian corvids. Family FRINGILLIDAE Subfamily CARDUELINAE Tribe DREPANIDINI The monophyly of the Drepanidini is supported by morphological and molec- ular data (Raikow 1977; Zusi 1978; Johnson et al. 1989), although one authority may still hold out for an independent origin of Melamprosops phaeosoma (Pratt 1979). In most cases we had to rely on phenetic resemblance to identify the new fossil taxa as members of this radiation. A few of the fossils were complete enough that derived osteological characters of the interorbital septurn and palatine pro- cess of the premaxilla could be cited as evidence of cardueline and hence pre- sumably drepanidine affinities. Even for the new species that are placed in the Drepanidini based on phenetics, we are satisfied that this assignment is correct NEW HAWAIIAN BIRDS 23 because each new taxon bears convincing resemblances to one or another branch of the drepanidine radiation. An independent test of our morphological assessment may be feasible if nucleotide sequences can be amplified from drepanidine fossils using the polymerase chain reaction (P•i•ibo 1989). Independent studies of myology (Raikow 1976), osteology (Zusi 1978), and DNA-DNA hybridization (Sibley and Ahlquist 1982; Bledsoe 1988) all indicate that the progenitor of the drepanidine radiation is likely to have been a cardueline finch. Although a recent study of allozymes favored a relationship between em- berizines and drepanidines (Johnson et al. 1989), the cardueline hypothesis is more highly corroborated, and we have assumed it to be correct. Consequently we place the finch-like taxa first in systematic order, and consider characters shared with carduelines to be primitive within the Drepanidini, although we have not attempted a phylogenetic analysis. The drepanidines as a group have been recognized by various authors as a distinct family (e.g., Amadon 1950), a subfamily (e.g., Pratt 1979), or a tribe (e.g., Olson and James 1982b; Sibley and Ahlquist 1982). Two subgroups of related species are usually recognized (in Pratt's scheme there are three). These subgroups are designated as subfamilies by Amadon, as tribes by Pratt. Neither of these options is available to us, because we continue to treat the entire radiation itself as a tribe. When we wish to distinguish the two subgroups, we therefore resort to the terminology of Perkins (1903), who was the first to group the predominantly nectarivorous red-and-black birds (Vestiaria, Drepanis, Hirnatione, Palmeria, and Ciridops), which he termed Division 1, apart from the rest of the radiation, termed Division 2. Our present taxonomic treatment of the drepanidines that retain finch-like bills departs from our earlier practice (Olson and James 1982b, 1984; James 1987; James et al. 1987) of lumping these birds into a single genus, Psittirostra sensu Amadon (1950), with species-groups treated as subgenera (Telespiza, Loxioides, Rhodacanthis, Chloridops, and Psittirostra). Instead, we have joined Pratt (1979), the American Ornithologists' Union (1983), and others in recognizing these taxa at the generic level. Arguments in favor of this change are that the genus Psittirostra sensu Areadon was united by what now appear to be primitive characters, and that it will make the magnitude of osteological differences between drepanidine genera more comparable to that between continental genera of Carduelinae. At a time when the relationships among the newly increased number of finch-billed species are poorly understood, this has the further advantage of avoiding the use of genera that have a high likelihood of being polyphyletic. In the future it may be advantageous to lump some of the genera recognized here, but when this is done it should be accompanied by evidence of monophyly (sensu Carroll 1987). Of the drepanidines that have departed from the finch-like bill shape, those with long, overhanging maxillary rostra (Psittirostra psittacea, Dysrnorodrepanis rnunroL Pseudonestor xanthophrys, Hernignathus lucidus, and Hernignathus wil- soni) receive little or no mention in our osteological comparisons because none of the new fossil taxa have this condition. The remainder of the drepanidines have thin bills that vary from short to very long, from straight o strongly decurved, and from sturdy to weak. We use the same generic taxonomy for these taxa as we have in previous papers (Olson and James 1982a, b, 1984), except that we have followed the commonly accepted practice of lumping Heterorhynchus with Hem- 24 ORNITHOLOGICAL MONOGRAPHS NO, 46 ignathus. This is a nomenclatural necessity in any case, because both genera have the same type species.. ' We'have made some minor revisions at the alpha level of taxonomy. The Kauai Amakihi (Loxops stejnegerO is listed as a full species rather than as a subspecies of Common Arnakihi (Loxops virens), as suggested by osteology and by allozyme data (Johnson et al. 1989). The Kauai Akepa is listed as a distinct species (Loxops caeruleirostris). The creepers from Lanai and Maui (Paroreomyza montana mon- tana .and P. montana newtoni) are no longer listed as subspecies of the Oahu Creeper (Paroreomyza maculata). These steps were suggested by Pratt (1979), and have been adopted by other authors in whole (Pratt et al. 1987) or in part (Berger 1981; American Ornithologists' Union 1983). Note that we do not accept Pratt's (1979) lumping of the nominal genera Viridonia (Greater Amakihi), Chlorodre- panis (Common and Kauai Amakihis), and Magumma (Akikiki) with Hemigna- thus, l•ut instead place .them with the akepas in Loxops. The pattern of differen- tiation in the individual island populations of akialoas is poorly known, and we have taken the conservative step of listing each named taxon as a species (Hemi- gnathus stejnegeri, H. lichtensteinL H. lanaiensis, and H. obscurus). Accurate identification of passefine fossils is frequently hindered by the great similarity between certain closely related species. This is true to a degree in the Drepanidini, in which morphological evolution of the postcranial skeleton has mainly affected stoutness and intramembral proportions of long bones, especially of the hindlimb. Most finch-billed taxa have stout tarsometatarsi that are relatively short in proportion to femur length, for example, while most thin-billed taxa have thin tarsometatarsi that are relatively long in proportion to femur length. Differ- ences of this sort may be useful for identifying associated skeletons, but usually are not sufficient o allow species-level identification of the isolated postcranial bones that are more frequently encountered in fossil deposits. On the other hand, the adaptive radiation of the drepanidine feeding apparatus has resulted in dramatic modifications ofosteology, such that even small fragments of maxillae and mandibles can often be identified accurately to species. Our descriptions of new drepanidine taxa rely almost entirely on characters of these elements, which preserve well as fossils and are abundantly represented in the Hawaiian deposits. We have furnished diagnoses based on bill osteology for the genera that contain new species. Characters of the maxilla used in our diagnoses, in addition to general bill shape, include the relative thickness of the dorsal and lateral nasal bars, modifications of the tomial crests, the dimensions and degree of definition of the narial openings, the contour of the posteroventral edge of the bone, the amount of lateral constriction, and the topography and degree of excavation of the ventral surface. Of these, the topography of the ventral surface and the contour of the posteroventral edge require further explanation. Finch-billed birds typically have the ventral surface of the rhamphotheca on the maxillary rostrum elaborated by some combination of grooves, ridges, troughs, horny plates, serrations, and stfiations. These structures have frequently been illustrated and correlated with feeding behavior and taxonomy (e.g., Sushkin 1924; Mayr et al. 1956; Bowman 1961; Ziswiler 1965). Although the rhamphothecal covering is missing from the fossil drepanidines, its variations are reflected to some degree in the underlying ventral surface of the bony maxilla, in which the NEW HAWAIIAN BIRDS 25 following structures can be observed in almost all drepanidines: a medial trough extending from the palatine articulations to the tip is bisected sagittally by a medial groove, and is bordered laterally by ventral ridges; small lateral troughs separate these ventral ridges from the tomia of the maxilla; the lateral troughs occlude with the posterior part of the mandibular tomia. Variations in the position and development of these structures are useful taxonomically. To assist the reader in comparing these structures in the photographic plates, their position in the finch-billed genera is identified diagrammatically in Fig. 9, which also illustrates variation in the contour of the posteroventral edge of the maxilla, ranging from irregular or straight through deeply V-shaped. Characters of the mandible used in the diagnoses include the degree of lateral constriction, the development of retroarticular processes, the relative size of the articular ends, the bluntness and curvature of the tomial crests, the shape of the middle part of the rami, particularly the relative depth at the angle of the mandible, the shape and thickness of the symphysis, and the shape and angle of the lateral cotylae. Comparative material examined.' We were able to examine at least partial skel- etal material of nearly all known species of Drepanidini, many of which are now extinct, so that bones had to be obtained from skins by the method outlined in Olson et al. (1987). The only taxa whose osteology was not examined were Hemi- gnathus lichtensteini and H. lanaiensis: Telespiza cantans males, USNM 289279, 289283, 499091, MVZ 148565; females, USNM 289280, 501414, 501648, 553311, MVZ 148564, PB 23537; unsexed, USNM 289281, 289282, 289284, 502155, 502223, AMNH 8882, 9060. T. ultima ma16, USNM 289278; females, USNM 289277, MVZ 124728, 124729, 124730. Loxioides bailleui males, MVZ 118827, 122620, 122621; probable females, USNM 560602, MVZ 118826, BBM-X 156555; unsexed, USNM 19098. ChloridøPs kona male, AMNH 453677 (bones removed from skin). Rhodacanthis palmeri male, AMNH 453623 (bones removed from skin). R. fiaviceps female, AMNH 453644 (bones removed from skin). Melam- prosops phaeosoma probable male, AMNH 810456 (bones removed from skin and from alcoholic trunk, AMNH 6371 [same individual]); unsexed, USNM 384745 (a nearly complete associated fossil skeleton from Auwahi Cave)2 Psitti- rostra psittacea male, MVZ 122619; unsexed, USNM 111.454 (bones removed from skin). Dysmorodrepanis munroi, BBM 4792 (bones removed fro m skin). Pseudonestor xanthophrys male, BMNH S/1961.11.46; female, BMNH S/ 1961.11.40. Hemignathus lucidus unsexed, BMNH S/1961 / 11.39. H. wilson ! male, MVZ 122610; female, MVZ 118830; unsexed, 122611. H. stejnegeri (•- H. pro- cents auct.) unsexed, USNM 19094, 19095 (bones, removed from alcoholic spec- imen). H. obscurus male, BBM 4434 (bones removed from skin); unsexed, BBM 109 (bones removed from skin). Oreomystis bairdi males, USNM 5'531'83, BBM 1231; unsexed, USNM 19096 (bones removed from alcoholic specimen'), 19097. Paroreomyza maculata (Oahu) unsexed, AMNH 5968. P. montana newtoni males, USNM 502187, 502188, 502189, MVZ 122615, 122616. P. fiammea unsexed, AMNH 453314 (bones removed from skin). Loxops eaeruleirostris males, USNM 553186, BBM 1229. L. coccineus coccineus male, 'MVZ 122613; unsexed, AMNH 9192. L. mana males, MVZ 122614, 118823. L. parvus males, USNM' 553187, 553188, 553189, 553191; females, USNM 502211, 553190; unsexed, USNM 19136. L. stejnegeri males, USNM 502195, 553192, 5531,94, 553196; females, 26 ORNITHOLOGICAL MONOGRAPHS NO. 46 3 1 Rf •• Mp Lb FiG. 9. Ventral maxillae of the finch-like genera ofdrepanidines, with Loxops virens for comparison. Bold lines illustrate the contour of the posteroventral margin of the maxilla, which can be straight (Oh), irregular (Ck), shallowly V-shaped (Rf, Tc, Lb), or more steeply V-shaped (Xf, Mp, Pp, and Lv, progressively). The following topographic features also vary in development among genera, as discussed in the text: 1, lateral trough; 2, ventral ridge; 3, medial groove; 4, medial trough. Oh, Orthiospiza howarthi; Rf, Rhodacanthis fiaviceps; Ck, Chloridops kona; Lv, Loxops virens; Tc, Telespiza cantans; Lb, Loxioides bailleui; Pp, Psittirostra psittacea; Mp, Melamprosops phaeosoma; Xf, Xestospiza fas- tigialis. USNM 502196, 553193, 553195; unsexed, USNM 502194. L. virensfiavus (= L. virens chloris auct.) females, USNM 553201, 553202, 553203, BBM-X 145538. L. v. wilsoni males, USNM 502191, 502192, 502193; female, USNM 502190. L. v. virens males, USNM 502197, 553209, 553210, 553213, 553214, MVZ 118755, 118760, 118766, 118767, 118782, 118786, 118787, 118788, PB 23868, 23869; NEW HAWAIIAN BIRDS 27 females, USNM 502198, 553211, MVZ 118756, 118761, 118762, 118763, 118765, 118772, 118814, PB 23870; unsexed, MVZ 118759, AMNI-I 9193, 9194, 9197. L. sagittirostris unsexed, AMN-H 453236 (bones removed from skin). Vestiaria coccinea males, USNM 502199, 502200, 502201, 502204, 553184, 553185, 553205, MVZ 118948, 118950, 118974, PB 23864, 23865; females, USNM 502202, 502203, MVZ 118942, 118949, 118955, PB 23867; unsexed, USNM 19130. Drepanisfunerea male, BBM 4712 (bones removed from skin). D. pacifica unsexed, BBM 2 (skull removed from skin). HirnationeJ•eethii unsexed, USNM 301141 (bones removed from skin), 301142 (bones removed from skin), 346246. H. sanguinea males, USNM 502205, 502206, 502207, 502209, 553182, 553217, 553218, MVZ 118858, 118883, 118885, 118886, 118927, PB 23861, 23862; females, USNM 502210, 553215, MVZ 118866, 118874, 118875, 118876, 118887, 118891; unsexed, USNM 19092, 553219, 502208. Palrneria dolei female, MVZ 122608. Ciridops anna unsexed, MCZ 19095 (bones removed from skin); BMNH 1939.12.9.58 (bones removed from alcoholic trunk specimen). Genus Telespiza Wilson, 1890 Fossil evidence has shown that the genus Telespiza was not always restricted in distribution to the Northwestern Hawaiian Islands but was widely distributed in the main chain of islands during the Holocene (Olson and James 1982b:39). Two new species from the main islands are described here, both smaller than the extant forms. In addition we have noted the existence of a possible third species, even smaller, of which we have only one mandibular ramus from Maul The four species of Telespiza hardly differ from each other in bill shape. Whereas other new drepanidines are diagnosed on qualitative osteological characters, the new species in this genus are diagnosed mainly on differences in bill size. To document size variation, we have included a longer list of paratypes and more osteometric data for this genus. We observed no overlap between any two species of Telespiza in the length of the maxilla measured from the lateral comer of the nasofrontal hinge, or the length of the mandible, although some overlap exists in other bill measurements (Tables 2 and 3, Fig. 10). Modem taxonomic practice would allow for allopatric populations that are differentiated mainly on size to be accommodated in one polytypic species. We recognize the new forms as full species because, even with the very patchy fossil record now available, three of the four taxa in the genus are known to have been sympatric with each other (Table 4). In the following generic diagnosis of Telespiza we have restricted our compar- isons to other finch-like drepanidine genera. Compared to Melarnprosops and Psittirostra, the narial opening is subcircular rather than oval, the dorsal nasal bar is broader and sturdier, the ventral surface of the maxilla is less excavated, the ventral ridges of the maxilla extend nearly to the tip rather than being confined posterolaterally, the posteroventral edge of the maxilla is less deeply V-shaped, and the mandibular tomium is more curved posteriorly. In addition, the maxillary rostrum does not overhang the mandibular rostrum in Telespiza to the same extent as in Psittirostra, and the lateral cotyla of the mandible is wider. Compared to Rhodacanthis, the bill is narrower, the dorsal nasal bar is broader and sturdier, the ventral ridges of the maxilla are not as prominent but are longer, extending nearly to the tip of the bill, and the medial trough of the maxilla is less excavated. 28 ORNITHOLOGICAL MONOGRAPHS NO. 46 TABLE 2 MEASUREMENTS (MM) OF THE MAXILLA IN Telespiza A_m) Loxioides. My_AN, 'STANDARD DEVIATION, AND RANGE ARE GIVEN FOR SAMPLE SIZES GREATER THAN 4; DATA ARE LISTED INDIVIDUALLY FOR SAMPLE SIZES OF 4 OR FEWER. H = HOLOTYPE Telespiza Telespiza Telespiza cantans Telespiza ultima persecutrix ypsilon Loxioides badleui Dorsal length 15.6 _+ 0.4 13.1 _+ 0.4 11.4 -+ 0.5 9.7 _+ 0.6 12.9 _+ 0.2 14.9-16.3 12.8-13.6 10.7-11.9 9.0-10.5 12.6-13.2 n=8 n=5 n=5 n=5 n=7 Ventral ength 9.0 -+ 0.4 7.1 _+ 0.3 6.8 -+ 0.6 5.3, 5.5 H, 6.2 _+ 0.3 8.1-9.5 6.8-7.6 6.1-7.6 5.5, 5.8 5.8-6.6 n=8 n=5 n=7 n=7 Length from lateral 15.5 + 0.4 13.4 -+ 0.3 11.8 -+ 0.4 9.4, 9.7 H, 12.8 --+ 0.2 comer ofnaso- 14.9-16.0 13.0-13.8 11.2-12.3 9.9, 10.5 12.5-13.2 frontal hinge n = 8 n = 5 n = 5 n = 7 Length from jugal 11.7 _+ 0.4 9.4 _+ 0.3 8.5 -+ 0.5 7.4 _+ 0.4 8.8 -+ 0.3 articulation 10.7-12.2 9.1-9.4 7.8-9.2 7.1-8.2 8.2-9.1 n=8 n=5 n=7 n=5 n=7 Length from 9.0 + 0.3 7.4 _+ 0.3 6.7 _+ 0.4 5.6 -+ 0.4 6.3 -+ 0.1 anterior narial 8.4-9.4 7.1-7.7 5.9-7.4 5.3-6.4 6.1-6.5 opening n = 8 n = 5 n = 7 n = 5 n = 7 Maximum width 8.2 _+ 0.4 6.9 _+ 0.3 6.4 _+ 0.3 5.5 -+ 0.2 8.1 _+ 0.2 7.8-8.8 6.6-7.3 6.1-7.1 5.3-5.7 7.9-8.4 n=8 n=5 n=9 n=5 n=6 Width of 9.1 _+ 0.5 7.3 +- 0.2 6.8 _+ 0.6 5.3, 5.5, 8.3 -+ 0.2 nasofrontal hinge 8.7-10.3 7.0-7.6 6.3-7.7 5.8 H 8.0-8.7 n=8 n=5 n=5 n=7 I_•ngth ofnarial 3.5 _+ 0.2 3.1 _+ 0.2 2.9 -+ 0.2 2.9 -+ 0.2 3.9 -+ 0.2 opening 3.1-3.7 3.0-3.4 2.5-3.1 2.7-3.1 3.6--4.2 n=8 n=5 n=9 n=7 n=6 Height ofnarial 3.4 + 0.2 2.8 + 0.1 2.6 + 0.1 2.4 + 0.2 3.5 + 0.2 oPening 3.2-3.7 2.7-2.9 2.4-2.8 2.2-2.7 3.1-3.7 n=8 n=5 n=8 n=7 n=6 Height through 7.3 + 0.4 6.3 + 0.3 5.1 + 0.3 4.0 + 0.4 6.7 + 0.3 lateral nasal bar 6.7-7.7 6.0-6.7 4.6-5.5 3.5--4.5 6.5-7.1 n=8 n=5 n=6 n=5 n=6 Minimum width of 2.4 + 0.2 2.0 + 0.1 1.9 + 0.2 1.5 + 0.1 2.3 + 0.2 dorsal nasal bar 2.2-2.7 1.8-2.2 1.7-2.2 1.3-1.6 2.0-2.6 n=8 n=5 n=8 n=7 n=7 Compared to Chloridops, the bill is less robust overall, the ventral ridges of the maxilla are longer but less prominent, and the tomial crest of the mandible is evenly decurved rather than nearly straight or ascending in the posterior portion. Telespiza is distinguished from Orthiospiza by the presence of a ventral trough of the maxilla, the much smaller narial openings, the shallowly V-shaped rather than straight posteroventral edge of the maxilla, the sturdier nasal bars, the less pointed mandible, and the less curved mandibular tomium. The bill of Telespiza is most similar to that of Loxioides, from which it differs by being longer and narrower rather than distinctly foreshortened, having a slightly wider lateral cotyla and a less abruptly decurved tomial crest of the mandible. These are relatively minor differences, so that it may be appropriate in the future to merge Telespiza NEW HAWAIIAN BIRDS 29 30 ORNITHOLOGICAL MONOGRAPHS NO. 46 TABLE 4 TH• DISTRIBUTION OF SPECIES IN THE GENUS Telespiza. INTRODUCED POPULATIONS ARE NOT INDICATED. X = EXTANT, • = FOSSIL Layman Nihoa Kauai Oahu Molokai Maui T. cantans X T. ultima X T. persecutrix T. ypsilon T. aft. ypsilon with Loxioides once again. For use of the spelling Telespiza as opposed to Tele- spyza, see Olson and James (1986). Telespiza persecutrix, new species (Figs. 11G-J, 12B, 13B) "Psittirostra (Telespyza), medium species" Olson and James, 1982b:39, 45; 1984:771; James, 1987:225. Holotype: Nearly complete maxilla lacking the dorsal nasal bar, with the ecteth- moids and part of the frontal attached, BPBM 158882 (Fig. 11G-I). Collected 23 July 1981 by A. Sinoto and P. C. McCoy. Type locality.' Site 50-Oa-B6-22, Barbers Point, Oahu, Hawaiian Islands. Distribution: Kauai: Makawehi Dunes. Oahu: Barbers Point and Ulupau Head. Etymology: Latin, persecutrix, one who follows pertinaciously; a sequel to the name T. ultima Bryan, 1917, given to the Nihoa Finch in the mistaken expectation that "the species under consideration is very liable to be the last native passefine bird t0be discovered in the Hawaiian Group" (Bryan 1917:70-71). Bryan's new finch was followed by the Nihoa Millerbird, Acrocephalus (familiaris) kingi (Wet- more, 1924), the drepanidines Dysmorodrepanis munroi Perkins, 1919 (collected in 1913), and Melamprosops phaeosoma Casey and Jacobi, 1974, and now by many new fossil species. The name is a feminine noun in apposition. Measurements (mm) ofholotype: Dorsal length, 11.4; ventral length, 6.4; length from lateral corner of nasofrontal hinge, 11.6; length from jugal articulation, 8.1; length from antefior nafial opening, 6.7; maximum ventral width, 6.7; width of nasofrontal hinge, 6.4; length of nafial opening, 2.5; height through lateral nasal bar, 5.3. Paratypes: Oahu: three complete maxillae, BPBM 177239, 177250 and 177254; maxffia with the cofused nasals broken posteriorly, BPBM 177240; maxilla lacking the left and part of the fight nasal bars, USNM 255230; maxilla lacking the left nasal bar, USNM 255562 (Fig. 11J); maxilla lacking the left nasal bar, BBM-X 155603; maxilla lacking the tip, BBM-X 153707; nearly intact juvenile maxilla, BBM-X 155885; mandible lacking part of the fight ramus, BPBM 177255; man- dible lacking part of the left ramus, BPBM 177158; mandible lacking parts of the fight ramus and left articular end, USNM 255208; two mandibular symphyses, USNM 255586 and BPBM 177194; mandible lacking part of the left ramus, USNM 447060 (Figs. 12B, 13B). Kauai: Two mandibles, each lacking part of the fight ramus, USNM 253888 and BBM-X 152693. NEW HAWAIIAN BIRDS 31 T. cantans n=• T. ultima n=5 T. persecutrix n=5 T. ypsilon n=4 T, cantans n=8 T. ultima n=5 T. persecutrix n=9 T. yps//on n=5 17 6 7 8 9 T. cantans n=8 T. u/t/ma n=5 T. persecutrix n=6 T. yps//on n:5 c [] ß 3 4 5 6 7 8 T. cantans n=8 D , T. ultima n=5 [] * T. persecutrix n=9 , • T. yps//on n=7 • ' 2.3 2.7 3.1 3.5 3.9 •. 10. Box plots oœmcasurcmc•ts (mm) oœtbc maxilla i• Telespiza. Tbc boxes J•dicat½ 25% of dc•otcs •c •c of •ucs, a•d ou•cn a• J•di•tcd by • astcds•. •, •c•½• from lateral co•cr of 32 ORNITHOLOGICAL MONOGRAPHS NO. 46 NEW HAWAIIAN BIRDS 33 ß A B FIG. 12. Mandibles of Telespiza in lateral view. A, T. ypsilon, new species (image is of the left side printed in reverse, USNM 254736); B, T. persecutrix, new species (USNM 447060); C, T. ultima, female (MVZ 124728): D. T. cantans. female (USNM 253711). Scale = I cm. Measurements ofparatypes: See Tables 2 and 3 and Fig. 10. Diagnosis: The bill is smaller than in T. cantans or T. ultima, but larger than in T. ypsilon, new species. Compared to T. ypsilon, the narial opening is slightly smaller in proportion to the overall size of the maxilla. Remarks: Of all the new species of drepanidines, this is the least differentiated in bill osteology. Nevertheless, the size differences between the three taxa are statistically significant. MANOVA F statistics comparing four measurements of the maxilla in T. ultima vs T. persecutrix and T. persecutrix vs T. ypsilon had a probability level of 0.01 (Wilks' Lambda), despite small sample sizes of four to five. The distribution of T. ultima, which is known from fossils to have occurred on islands to the east and west of T. persecutrix, argues that these two taxa are not subspecies. It is at least as plausible that T. ultima formerly had a continuous distribution and was thus sympatric with T. persecutrix. Recognizing T. persecutrix as a distinct species serves to emphasize that mem- bers of this genus were evidently widely distributed and that a high degree of sympatry existed among them. It is also consistent with the modern reversal of opinion regarding T. ultima, which was long treated as a subspecies of T. cantans but is now considered to be a distinct species because of differences in size, FIG. 11. Maxillae of Telesptza in three views. A, lateral, B, dorsal, and C, ventral views of T. cantans, female (MVZ 148564); D, lateral, E, dorsal, and F, ventral views of T. ultima, female (MVZ 124728); G, lateral, H, dorsal, and I, ventral views of T. persecutrix, new species, holotype (BPBM 158882); J. dorsal view of T. persecutrix (USNM 255562); K. lateral. L, dorsal, and M, ventral views of T. ypsilon, new species, holotype (USNM 253711). Scale = I cm. 34 ORNITHOLOGICAL MONOGRAPHS NO 46 B D FIG. 13. Mandibles of Telespiza in dorsal view. A, T. ypsdon, new species (USNM 254736); B. T. persecutrix, new species (447060); C, T. ultima, female (MVZ 124728); D, T. cantans, female (USNM 253711). Scale = I cm. plumage, and plumage sequence (Banks and Laybourne 1977), as well as by its fossil sympatry with T. cantans on Molokai (Olson and James 1982b). All of the named species of Telespiza occur on more than one island, and three of the four were found together on Molokai (Table 4). As the fossil record improves, we might expect to find three or more species on each of the main islands. NEW HAWAIIAN BIRDS 35 Telespiga ypsilon, new species (Figs. 11 K-M, 12A, 13A) "Psittirostra (Telespyza), small species" Olson and James, 1982b:39, 45; 1984:772. Holotype: Complete maxilla, USNM 253711 (Fig. 11 K-M). Collected 12 or 19 July 1976 by Storrs L. Olson and Joan Aftdem. Type locality: Site 20, Ilio Point, Molokai, Hawaiian Islands. Distribution: Molokai: Ilio Point. Maui: Puu Naio Cave. Etymology: Greek, ypsilon, twentieth letter of the Greek alphabet, in reference to the number of the type locality. The name is a neuter noun in apposition. Measurements (ram) ofholotype: Dorsal length, 9.3; length from jugal articu- lation, 7.3; length from anterior narial opening, 5.3; maximum ventral width, 5.5; length of narial opening, 2.9; height of narial opening, 2.2; height through lateral nasal bar, 4.1; minimum width of dorsal nasal bar, 1.3. For additional measure- ments, see Table 2. Paratypes: Molokai, Ilio Point: complete maxilla with tip slightly abraded, USNM 253712; nearly complete maxilla lacking tip, USNM 254698; maxilla lacking right nasal bar and ventral rim of right narial opening, USNM 254850; maxilla lacking right nasal bar, USNM 445779; maxilla lacking tip and left nasal bar, BPBM 178140; maxilla lacking right nasal bar, BPBM 178141; mandible lacking parts of the symphysis and right ramus, USNM 254736 (Figs. 12A, 13A); mandible lacking parts of both rami, USNM 255013. Maui, Puu Naio Cave: The portion of a maxilla anterior to the narial openings, USNM 445780; mandibular symphysis, USNM 445781. Measurements ofparatypes: See Tables 2 and 3 and Fig. 10. Diagnosis: The bill is smaller overall, and the narial openings are somewhat enlarged relative to the size of the maxilla, compared to Telespiza cantans, T. ultima, or T. persecutrix. Remarks: T. ypsilon was by far the commonest passeme prey item of the long- legged owl, Grallistrix geleches, in the pellet deposits at Ilio Point on Molokai. One hundred and thirty-one pieces of the bill of this species were identified from Sites 16 and 20, making up 53% of the minimum number of individual drepan- idines, which suggests that T. ypsilon was fairly abundant in this harsh, arid environment. The species was not restricted to extremely arid coastal habitats, however, as the specimens from Puu Naio Cave on Maui originated at about 305 m elevation in a region of more moderate climate. Telespiza aft. ypsilon, Maui Material.' A left mandibular ramus, USNM 445782. Distribution: East Maui: Lua Lepo. Measurements: See Table 3. Remarks: This specimen is 22% smaller than the next larger specimen of Tele- spiza (T. ypsilon), based on the average difference between seven measurements compared in Table 3. In all likelihood it represents a distinct species. The specimen is also notable as an elevational record, because it was found at a much higher elevation (808 m) than any locality where Telespiza has previously been reported. Besides this record and the few specimens of T. ypsilon from Puu Naio Cave (305 m), the genus is known only from localities near sea level. 36 ORNITHOLOGICAL MONOGRAPHS NO. 46 Genus Chloridops Wilson, 1888 In Chloridops the bill is robust and heavily ossified, to an extent comparable to continental grosbeaks. The thick walls of the maxilla and mandibular symphysis encase a dense network of uncommonly sturdy bony trabeculae that presumably functioned to disperse the strong forces generated when cracking hard seeds with the bill. The relatively straight tomial crest of the maxilla, with a slight to marked ventral bulge about one-third of the distance from the tip, distinguishes Chloridops from all related genera, in which the tomial crest is evenly dectawed. Prominent but blunt ventral ridges of the maxilla are also unique to Chloridops. These ridges are prominent but sharp in Rhodacanthis, and are more weakly developed in all other drepanidines. In Chloridops, the narial opening is never elongate antero- posteriorly, but usually resembles Telespiza, Loxioides, and Rhodacanthis in being subcircular, while in C. regiskongi, new species, the narial opening is markedly higher than it is long. The mandible in Chloridops is extremely robust, with the walls of the bony symphysis thicker, the ramus deeper in relation to its length, and the articular end enlarged, compared to other drepanidines. The anterior edge of the symphysis is broadly curved rather than pointed, a condition also found in Loxioides. The posterior third of the tomial crest of the mandible is blunt or flattened to a greater degree than in other drepanidines. Chloridops sp., Kauai "Psittirostra (Chloridops), Kauai species" Olson and James, 1982b:40, 44. "Psitirostra [sic] (Chloridops), Kauai sp." Olson and James, 1984:771. Material.' Symphysis and left ramus of a damaged mandible, USNM 253929; the middle part of a right ramus, USNM 254980; right ramus without symphysis, USNM 253904; two fragments of palatines, USNM 254988 and 254989; two right quadrates, USNM 254981 and 445783; left quadrate, USNM 254982. Distribution: Kauai: Makawehi Dunes. Measurements: See Table 6. Remarks: The mandible is similar to that of Chloridops wahl new species, except that the lingual walls of the mandibular symphysis are more widely set apart, creating a broader lingual trough. Although this difference between C. wahl and the mandible from Kauai is quite striking, a larger sample is needed to evaluate how much variation may occur within species. It is not unlikely that the Kauai Chloridops is a distinct species, because intraspecific variation of this magnitude is not apparent in the series of mandibles of C. regiskong•, new species. In any case, the narrower lingual trough is present only in C. wahl and C. kona; the trough is wider in other forms of Chloridops. Chloridops wahl, new species (Figs. 14A, 15C, D, 16A) "Psittirostra (Chloridops), lesser Oahu species" (Olson and James, 1982b: 40, 45; 1984: 771; James, 1987:225, 228. Holotype: Nearly intact maxilla lacking a small part of the left nasal, BBM-X 155524 (Figs. 14A, 16A). Collected July or August 1977 by Aki Sinoto. Type locality.' Site 50-Oa-B6-100B, Barbers Point, Oahu, Hawaiian Islands. Distribution: Oahu: Barbers Point and Ulupau Head. Maui: Puu Naio Cave. NEW HAWAIIAN BIRDS 37 A B FIG. 14. Maxillae of Chloridops in dorsal (left) and ventral (right) views. A, C. wahl, new species, holotype (BBM-X 155524); B, C. kona, male (AMNH 453677); C, C. regiskongi, new species (dorsal view is ofholotype, BPBM 158742; ventral view is of BPBM 158814). Scale = I cm. Etymology: Hawaiian, wahl to cleave or break in pieces; in reference to the function of the bill in cracking hard seeds. Measurements ofholotype: See Table 5. Paratypes: Oahu, Barbers Point: maxilla lacking part of nasals, USNM 255568; nearly intact maxilla, BBM-X 155155; articular end of mandible lacking medial A ,,• B C / D •-?.._._ ',-,r,• • NEW HAWAIIAN BIRDS 39 TABLE 5 MEASUREMENTS (MM) OF THE MAXILLA IN Chloridops. TI•œ MFa•S•M•S IN P/mENTI•SES AgE ESTIMATES DUE TO THE 1VIISSING TIP OF THE 1VIAXILLA IN THE HOLOTYPE OF C. regiskongi. H = HOLOTYPE Chloridops Chloridops Chloridops Chloridops wahl, Oahu wahl, Maul kona regiskongi Dorsal length Ventral length Length from lateral comer of nasofrontal hinge Length from jugal articulation Length from anterior narial opening Maximum width Width of nasofrontal hinge Length of narial opening Height of narial opening Height through lateral nasal bar Minimum width of dorsal nasal bar 18.4 H 18.5 22.2 (25.5 H) 11.7 H 12.0 12.8 (14.9 H) 18.8 H 18.9 22.1 (25.2 H) 14.4 a 14.0 16.0 (18.6 H) 10.2 a 9.7 11.8 (16.2 H) 12.2 H, 12.4, 12.5 12.5 13.2 14.9 a 12.3 -- 12.3 16.2 a 3.9 H, 4.0, 4.4 4.3 4.6 3.6 a 3.8 H, 3.8, 3.9 4.3 4.4 5.5 a 7.9, 8.5 H 9.1 10.7 13.7 H 2.5, 2.7, 3.1 H 2.5 2.2 6.2 H process, USNM 445785; mandibular symphysis with left ramus, BBM-X 151278 (Figs. 15D, 16A); left mandibular ramus lacking the medial process and most of the symphysis, BPBM 158681 (Fig. 15C). Maul, Puu Naio Cave: maxilla lacking part of the left nasal, USNM 445784. Measurements ofparatypes: See Tables 5 and 6. Diagnosis: Similar to Chloridops kona but with the bill smaller and shorter, and the dorsal nasal bar broader anteriorly. Chloridops wahl and C. kona are very similar in other characters of bill osteology, including the lateral cotyla which is elevated and angled posteromedially, the narrow sagittal groove in the lingual surface of the mandibular symphysis, the flattened posterior portion of the tomial TABLE 6 MEASUREMENTS (MM) OF THE MANDIBLE IN Chloridops Chlori- Chlori- dops sp., Chloridops dops sp., Chlori- Kauai wahl. Oahu Maul dops kona Chloridops regiskongi Total length 27.9 -- 21.2 31.1 36.5, 37.3 Length oftomial crest 14.3 14.0 11.0 16.0 17.1, 17.8, 19.9 Symphysis length 9.2 10.0 6.8 12.4 10.9, 11.1, 12.6 Greatest width of symphysis 13.1 -- 10.1 14.7 16.5, 17.8, 18.0 Greatest width of mandible -- -- 16.8 23.2 28.8, 29.0 Symphysis height 5.7 6.2 4.5 7.1 8.5, 9.4 Length of lateral cotyla 4.3 -- 3.2 4.3 5.6, 5.6, 6.3 Width of articular end with medial process 7.1 -- 5.4 -- -- Height at lateral cotyla 3.6 4.1, 4.2 3.0 4.4 4.9, 5.0, 5.2, 6.0 Height at angle of mandible 9.7 8.6, 9.4 6.5 10.9 10.0, 10.9, 11.6 Ramus length (middle part) 12.5 11.7, 12.0 10.0 14.4 14.2, 14.2, 14.4, 14.9 FIG. 15. Mandibles of Chloridops in dorsal view. A, C. regiskongi, new species (USNM 445789); B, C. kona, male (AMNH 435677); C, C. wahl, new species (BPBM 158681); D, C. wahl (BBM-X 151278). Scale = 1 cm. 40 ORNITHOLOGICAL MONOGRAPHS NO. 46 crest of the mandible, the very blunt, close-set ventral ridges of the maxilla, and the presence of a foramen on each nasal above the narial opening. The mandible of C. wahi also resembles those referred to Chloridops sp. from Maui and Kauai, except that it has a narrower lingual trough with a deeper sagittal groove in the symphysis, and is larger than the spedmen of Chloridops sp. from Maul The bill differs from that of C. regiskongi, new species, in having the maxilla less deep, the blunt sagittal crest of the maxilla absent, the dorsal nasal bar much narrower, the narial openings subcircular, the lateral nasal bars more robust, the nasal septum usually partly ossified, the ventral ridges of the maxilla more blunt and close-set, the antorbital space relatively small, the posterior margin of the mandibular symphysis nearly straight, the lateral cotylae of the mandible elevated and angled posteromedially; in the presence of a foramen above each narial opening, and in the presence of a sagittal groove in the mandibular symphysis. Remarks: Although the bill differs in qualitative characters and in size, this species is clearly allied to C. kona and to the spedmens from Kauai and Maui referred only to Chloridops sp. The only skeletal element that is known from all of these forms is the mandible, so that the sagittal groove in the mandibular symphysis is the only synapomorphy yet detected in all of them. This character is more pronounced in C. kona and C. wahi than in the specimens referred to Chloridops sp. Chloridops sp., Maui Material: A complete mandible, the left ramus having been broken and repaired, USNM 445786. Distribution: Maui: Puu Naio Cave. Measurements: See Table 6. Remarks: This almost perfectly preserved mandible is the smallest existing specimen referable to Chloridops. The next larger mandible, a paratype of C. wahi from Oahu (BBM-X 151278), measures 18% larger in estimated length. The small mandible from Maui agrees with C. wahi and C. kona in the characters that set these species apart from C. regiskongi (see diagnoses ofC. wahi and C. regiskongi), and differs from C. wahi and C. kona in having the walls of the symphysis more widely spaced. Additional specimens of Chloridops are needed from Maui before we can decide whether to interpret this mandible as an extreme of intraspecific variation in C. wahi or as a distinct species. Chloridops regiskongi, new species (Figs. 14C, 15A, 16A-B) "Psittirostra (Chloridops), giant Oahu species" Olson and James, 1982b:40, 45; 1984: 771; James, 1987:225, 228. Holotype: Maxilla lacking the tip and the right tomial crest, with a small hole anterior to the left narial opening, BPBM 158742 (Figs. 14C, 17A). Collected 14 August 1981 by Aki Sinoto, Patrick C. McCoy and others. Type locality.' Site 50-Oa-B6-22, Barbers Point, Oahu, Hawaiian Islands. Distribution: Oahu: Barbers Point and Ulupau Head. Etymology: "Of King Kong" from the Latin genitive of rex, king, plus Kong; NEW HAWAIIAN BIRDS 41 in reference to the great size of this finch by allusion to the giant primate of cinema fame. In a newspaper article, Olson was once quoted as saying this species was "a giant, gargantuan, a King Kong finch" (Benson ! 977), an appellation that would never have occurred to him, this being a typical example of the liberties taken with quotation marks by the print media. Nevertheless, because of its ridicu- lousness, the name naturally stuck and we found ourselves referring jocularly to this species as the "King Kong finch" ever afterwards. Measurements of holotype: See Table 5. Paratypes (all from Barbers Point): Maxilla with only the ventral surface and tomial crests remaining, BPBM ! 588 ! 4 (Figs. ! 4C, 17C); nearly intact mandible lacking part of both roedial processes, USNM 445789 (Figs. 15B, 17B); mandible lacking part of both rami and roedial processes, USNM 445788; mandible lacking the posterior half of the right ramus and left roedial process, USNM 445787; mandible lacking part of the symphysis and left ramus, BBM-X 151419; left quadrate, USNM 445790. Measurements ofparatypes: See Table 6. Diagnosis: A grosbeak finch possessing the largest bill known among finch-like drepanidines. The bill differs markedly from the other two species of Chloridops in shape as well. The maxilla is much deeper and is sharply pointed, the sides of the maxilla rising steeply to form a blunt crest that extends along the dorsal midline from the tip through the dorsal nasal bar. The dorsal nasal bar is extremely broad compared to other drepanidines, so broad as to occlude the anterior part of the narial openings, which are consequently higher than long. The dorsal nasal bar is also distinctively elevated at its posterior margin. C. regiskongi differs further from C. kona and C. wahi as follows: the lateral nasal bars are weaker; the nasal septurn is unossified (usually partly ossified in C. kona and C. wahi); the ventral surface of the maxilla does not extend as far posteriorly, so that there is no ossified floor beneath the narial openings, which, with the extreme depth of the maxilla and the fairly extensive area of co-ossified nasals, creates a greater antorbital space; the prominent ventral ridges of the maxilla are less blunt and less close- set, tending more toward the configuration in Rhodacanthis; the foramina above the narial openings are absent; the mandibular symphysis lacks a distinct sagittal groove; in ventral aspect, the posterior margin of the symphysis is curved, as is more typical of finches, rather than nearly straight; and the lateral cotylae lie approximately parallel to each other and are not elevated, as opposed to being elevated and angled posteromedially. Remarks: In addition to the character differences mentioned in the diagnosis, the bill of C. regiskongi appears to have developed along a different allometric trajectory from the other species in the genus, as it differs further from C. kona and C. wahi, but resembles the smallest known mandible of the genus (Chloridops sp., Maul) in being less extensively ossified, with the posterior portion of the tomial crest of the mandible not as broad and flat, the middle part of the ramus a little less deep (and consequently the angle of the mandible somewhat less acute), and the surface for attachment of M. depressor mandibulae less expanded. Although we have placed this species in the genus Chloridops, its osteological differences from other members of the genus are extensive enough to raise the possibility of parallel evolution in two lineages from weaker-billed finches. In particular, the broad, parallel lateral cotylae of the mandible combined with the 42 ORNITHOLOGICAL MONOGRAPHS NO. 46 NEW HAWAIIAN BIRDS 43 A F•G. 17. Maxillae and mandible of Chloridops regiskongi, new species, in lateral view. A, maxilla, holotype (BPBM 158742); B, mandible CLISNM 445789); C, maxilla lacking the dorsal portion (BPBM 158814). Scale = 1 cm. prominent ventral ridges and weak lateral nasal bars of the maxilla could be taken as evidence that C. regiskongi is a sister taxon of Rhodacanthis. Genus Incertae Sedis, Unassigned Maui Finch "P[sittirostra] (Chloridops) sp." James et al., 1987:2353. Material: A fragment of cranium including the frontal and parts of the inter- oribital septurn and maxilla, USNM 445791. Distribution: Maui: Puu Naio Cave. Measurements (mm): Interorbital width of frontal, 9.8; minimum width of dorsal nasal bar, 2.4; height of narial opening, 4.3; length of narial opening, 4.5. Remarks: This provocative fragment preserves just enough morphology to make it clear that it is something new, but not enough to determine its relationships. It represents a finch about the size of C. wahl, with a sturdy, arched maxilla. Although we had earlier referred it to Chloridops, it differs from C. kona and C. wahi in having the interorbital septurn less thickened, the lateral nasal bars as FIG. 16. Maxillae and mandibles of Chloridops wahl new species, and C. kona, in lateral view, oriented approximately as in life. A, C. wahi (maxilla = holotype BBM-X 155524, mandible = BBM-X 151278; both images are of the left side printed in reverse); B, C. kona male (AMNH 453677). Scale = lcm. 44 ORNITHOLOGICAL MONOGRAPHS NO. 46 well as the jugal and palatine articulations of the mandible weaker, and the lateral troughs shallower. It differs from C. regiskongi n having subcircular narial open- ings and a much smaller antorbital space. It differs from Rhodacanthis in having the medial and lateral troughs of the maxilla shallower, from Orthiospiza in having the narial openings smaller and the tomial crest broader posteriorly, and from Telespiza in not having the posteroventral edge of the maxilla distinctly V-shaped. Little more can be said about this bird until better specimens are found. It is worth mentioning that this fragment is not the only enigmatic bit of finch bill in the USNM collections from Maul. Fossils we have examined strongly suggest hat at least two more finch-billed species exclusive of Telespiza will eventually be described from Maul, a prospect that should provide some impetus for an inten- sified fossil search. Genus Rhodacanthis Rothschild, 1892 The two extinct finches in this genus, Rhodacanthispalmeri (Greater Koa Finch) and R. fiaviceps (Lesser Koa Finch), were restricted to the island of Hawaii when they were discovered by late 19th-century collectors. As no skeletons were pre- served of either species, we removed the bones from one skin specimen each of R. palmeri and R. fiaviceps. These forms had sometimes been considered con- specific, although we found qualitative differences in bill osteology that support Amadon's (1950) view of them as distinct, sympatric species. Characters diagnosing Rhodacanthis are as follows. The ventral trough of the maxilla is deeper than in other finch-billed drepanidines. The ventral ridges are more prominent than in other genera except Chloridops, and are sharp, whereas in Chloridops they are blunt. As in Chloridops, the ventral ridges extend about three-fourths of the distance to the tip of the maxilla, as opposed to nearly reaching the tip as in Telespiza, Loxioides, and Orthiospiza, new genus, or being confined posterolaterally as in Psittirostra and Melamprosops. The narial openings are subcircular, not elongated as in Psittirostra and Melamprosops, and not greatly enlarged as in Orthiospiza. The dorsal nasal bar is stouter than in Orthiospiza, Melamprosops, and Psittirostra but less so than in Chloridops, Telespiza, and Loxioides. The mandible in Rhodacanthis is sturdy and deep compared to Te- lespiza, Loxioides, Psittirostra, and Melamprosops. It is distinctly pointed rather than having a broad tip like Chloridops or Loxioides. The symphysis is longer than in Orthiospiza and Loxioides. The lateral cotylae of the mandible can be distinguished from other drepanidines, except Chloridops regiskongi, through the combination of not being elevated, being very broad in the middle, and being angled slightly in the posterolateral rather than posteromedial direction. We have recorded fossils of Rhodacanthis from Oahu (Olson and James 1982b; James 1987) and Maul (James et al. 1987). These differ from R. palmeri and R. fiaviceps, but not in ways that we consider important enough to justify formally describing new species at this time, considering that our comparative material is limited to only one partial skeleton of each species. Rhodacanthis aft. palmeri Rothschild, 1892 P[sittirostra] (Rhodacanthis) aft. palrneri James et al., 1987:2353. Material: Maxilla with the dorsal part missing from the posterior half, USNM 445792. NEW HAWAIIAN BIRDS 45 TABLE 7 MEASUREMENTS (MM) OF THE iVIAXILLA IN Rhodacanthis AND Orthiospiza. H = HOLOTYPE Rhoda- Rh•da- Rh•da- canthis Rhodao canthis Orthio- canthis aft. canthis aft. spiza paltoefl paltoefl fiaviceps fiaviceps howarthi Dorsal length 20.5 -- 17.7 17.3 15.3 h**, 17.4 Ventral length 13.8 -- 10.9 11.0, 11.1, 11.2 9.7 H*, 11.7 Length from lateral corner of nasofrontal hinge 20.4 -- 17.7 17.8 16.0 H*, 17.5 Length from jugal articulation 15.5 15.7 13.4 13.0, 13.1, 13.5 11.9 H*, 12.4 Length from anterior narial opening 12.3 -- 9.8 10.2, 9.7, 9.6 6.5 •*, 9.3 Width of nasofrontal hinge 11.7 -- 10.6 11.5 9.3, 9.4 • Length of narial opening 4.5 -- 4.5 4.5, 4.7, 5.2 5.4, 5.9 • Height of narial opening 4.6 -- 3.5 4.0, 4.0, 4.1 4.8 •, 4.9 Height through lateral nasal bar 8.6 -- 8.3 8.4 8.4 H, 9.2 Minimum width of dorsal nasal bar 2.0 -- 1.7 1.7, 1.8, 1.6 1.4, 1.6 H * Approximately 2 mm should be added to these measurements to compensate for the broken tip of the maxilla in the holotype of O. howarthi. Distribution: Maui: Puu Naio Cave. (R. palmeri is known historically from the island of Hawaii.) Measurements: See Tables 7 and 8. Remarks: This maxilla differs from R. fiaviceps and agrees with R. palmeri in being larger and having a deeper, wider ventral trough of the maxilla. The trough is even deeper but narrower than in the comparative specimen ofR. palmeri, yet it may fall within the range of variation for that species. Rhodacanthis aft. fiaviceps Rothschild, 1892 "Psittirostra (Rhodacanthis) fiaviceps" Olson and James, 1982b:39, 45. "Psittirostra (Rhodacanthis) sp." James, 1987:225, 228. " P[ sittirostra] (Rhodacanthis) aff. fiaviceps" James et al., 1987:2353. TABLE 8 MV_•SUm•M•NTS (MM) OF •-Iœ MANDIBLE IN Rhodacanthis a•D Orthiospiza. H = HOLOTYPE Rhodacanthis Rhodacanthis Rhodacanthis Orthiospiza palmeri fiaviceps aft. fiaviceps howarthi Total length 30.0 26.1 26.3 26.6 Length of tomial crest 16.5 12.9 13.0, 13.7 12.4 Symphysis length 10.3 8.7 7.4, 7.9 7.1 Greatest width of symphysis 12.4 11.4 11.4, 11.6 11.1 Greatest width of mandible 20.1 18.8 -- 19.3 Symphysis height 6.0 5.7 4.6, 4.8 6.1 Width of articular end with roedial process 6.0 -- 5.4 6.4, 6.9 H Height at lateral cotyla 2.8 3.1 2.8 2.5, 2.7 • Height at angle of mandible 6.2 6.2 5.8 5.9, 7.8 • Ramus length (middle part) 12.5 11.0 11.1 12.6, 13.4 46 ORNITHOLOGICAL MONOGRAPHS NO. 46 Material.' Puu Naio Cave, Maui: Maxilla lacking left nasal bar, USNM 445794; mandible lacking posterior third of left ramus, USNM 445793; mandibular sym- physis, USNM 445796. Barbers Point, Oahu: Maxilla, BPBM 158861. Ulupau Head, Oahu: Maxilla lacking parts of the fused nasals, USNM 445795. Distribution: Oahu: Barbers Point and Ulupau Head. Maui: Puu Naio Cave. (R. fiaviceps is known historically from the island of Hawaii.) Measurements: See Tables 7 and 8. Remarks: These specimens are similar in size to R. fiaviceps but differ in having a larger narial opening, a deeper medial trough on the ventral surface of the maxilla, and a decurved rather than straight anterior portion of the tomial crest of the mandible. In the last two characters, the fossils agree more with the larger R. palmeri than with R. fiaviceps. Faced with such an ambiguous combination of characters, we are unable to suggest whether this form is a distinct species, is conspecific with R. fiaviceps, or is conspecific with R. palmeri. Orthiospiza, new genus Type species: Orthiospiza howarthL new species. Included species: Type-species only. Distribution: Maui: higher elevations on the south and east slopes of Mr. Ha- leakala. Etymology: Greek, orthios, high, lofty, and spiza, a finch, to emphasize that specimens of this genus have so far been found only at relatively high elevations. The gender is feminine. Diagnosis: Large drepanidines with arched, finch-like bills, distinguished by the greatly enlarged narial openings, the absence of a ventral trough on the maxilla, and the straight rather than indented posteroventral contour of the maxilla. The cranial fenestra is smaller than in any other finch-billed drepanidine in which the cranium is known, except Chloridops kona. The muscle scars on the cranium and mandible reflect a well-developed musculature for adduction of the mandible, with robust zygoraatic processes, prominent interorbital cristae, a distinct crest delimiting the main area of attachment of adductor muscles on the skull, and partly ossified aponeuroses preserved on the coronoid processes and on the ventral margins of the mandibular foramina in the adult specimen. By contrast, the palatines are underdeveloped, with the transpalatine processes short and weak and the prepalatine bars very weak compared to Rhodacanthis and other large finches. The prepalatine bars also differ from the relatively thin bars of Psittirostra in being very short rather than elongated. The surfaces on the maxilla for artic- ulation of the palatines and jugals are distinctly weaker than expected for a finch with such a sturdy bill. The maxilla is broad and pointed, with an arched dorsal surface. The tomial crest of the maxilla is evenly decurved, as in Telespiza, and the dorsal nasal bar is elongated, as in Psittirostra. Broad, moderately excavated lateral grooves pro- vide the only relief on the otherwise nearly flat ventral surface of the maxilla. The ventral ridges are very close-set, as in Chloridops kona, but extend nearly to the tip of the maxilla, as in Telespiza and Loxioides. The ventral ridges differ from other genera in being not at all prominent, and in being rugose in the adult. NEW HAWAIIAN BIRDS 47 A C D E FtG. 18. The cranium, palatine, and quadrate of the holotype of Orthiospiza howarthi. new genus and species (BPBM 160712). A, posterior view of the interior of the brain case; B, the same fragment in anterior view, showing the prominent interorbital cristae, the relatively small cranial fenestrae, and trabeculae along the midline indicating where the interorbital septurn is broken away; C, part of the fused palatines with the right prepalatine bar and transpalatine process; D, fragment of the braincase with the zygomatic process; E, left quadrate. Scale = I cm. Between these ridges, where the medial trough usually occurs, the maxilla is flat and is also rugose in the adult. The mandible is deep and pointed as in Rhodacanthis, but has a short symphysis and a strongly and evenly decurved tomial crest as in Loxioides. The shape of the lateral cotylae, which are elevated, angled posteromedially, and have the anterior portion not much wider than the posterior portion, is reminiscent of the much smaller cotylae of Xestospiza fastigialis, new genus and species. Orthlosl)i•a howarthi, new species (Figs. 18, 19D-G, 20A-B, D-E, 21) Holotype: Partial associated skeleton, BPBM 160712. Collected between 24 February and 4 March 1984 by Francis G. Howarth, Fred D. Stone, Betsy H. Gagng and others. The specimen consists of the maxilla lacking the tip (Fig. 19D, F, G), fragments of the cranium preserving the posterior wall of the orbits (Fig. 18A, B), the basitemporal plate, the left zygomatic process (Fig. 18D), and the supraoccipital, the right pterygoid, the fused palatines with the left side mostly 48 ORNITHOLOGICAL MONOGRAPHS NO. 46 FIG. 19. Maxillae of Rhodacanthisfiaviceps and Orthiospiza howarthi, new genus and species. A, lateral, B, ventral, C, dorsal views ofR. fiaviceps female {AMNH 453644); D, lateral, F, ventral, G, dorsal views of O. howarthi holotype (BPBM 160712; lateral view is of the left side printed in reverse): E. ventral view ofO. howarthi, immature (USNM 370557). Scale = I cm. missing (Fig. 18C), both quadrates (Fig. 18E), both rami of the mandible but not the symphysis (Fig. 20B, D), the axis and eight additional cervical vertebrae, three thoracic vertebrae, two caudal vertebrae, the pelvis (Fig. 21A), the anterior half of the fight scapula, the dorsal half of the fight coracoid, the ventral third of the left coracoid, both humeri (each missing parts of both ends; Fig. 21B), the right ulna (Fig. 21C), fragments of the shaft of the left ulna, both carpometacarpi with NEW HAWAIIAN BIRDS 49 damaged proximal ends (Fig. 21D), both femora with damaged distal ends, the right tibiotarsus with damaged ends, the broken left tibiotarsus, proximal ends with parts of the shafts of both tarsometatarsi, the damaged distal end of the left tarsometatarsus, three pedal phalanges, and some unidentified bone fragments. Type locality: Lua Manu (1,830 m), Kipahulu Valley, Maul, Hawaiian Islands. The location of the holotype within the cave coresponds with "Site L" on the cave map drawn by Fred Stone and Francis Howarth, a copy of which is on file with the accession record at the Bishop Museum (BPBM Accession 1984.116). Distribution: Maul: Lua Manu, Puu Makua Cave, and Auwahi Cave. Etymology: To Francis G. Howarth, for his studies of the entomology of Ha- waiian caves and for his attentivehess to potential fossil deposits in lava tubes. Measurements (ram) of holotype: The comparable measurement of Rhodacan- this palmeri (AMNH 453623) is given in parentheses when available. Humerus: length (estimated), 25.0 (24.5); proximal width, 7.2 (7.8); mid-shaft width, 2.4 (2.4). Ulna: length, 29.8. Carpometacarpus: length, 16.2. Synsacrum: length, 2.5. Pelvis: width between antitrochanters, 19.1. Femur: length, 26.9; proximal width, 4.8; mid-shaft depth, 1.9. Tibiotarsus: length without cnemial crest, 41.5. Tar- sometatarsus: proximal width, 4.3 (4.2); mid-shaft width, 1.7 (1.7). For additional measurements of holotype, see Tables 7 and 8. Paratypes: Auwahi Cave: Maxilla lacking lateral nasal bars, USNM 370557 (Fig. 19E); complete mandible, USNM 372839 (Fig. 20A, E); humerus lacking the head and distal condyles, USNM 372840. These bones appear to be from a single juvenile individual. Puu Makua Cave: Complete maxilla, not fully mature, USNM 445797; man- dible in two pieces, lacking the anterior two-thirds of the right ramus, not fully mature, USNM 445798. These bones may be from a single individual. Measurements ofparatypes: See Tables 7 and 8. Diagnosis: As for the genus. Remarks: The body size is comparable to Rhodacanthis palmeri, the largest of the finch-billed drepanidines for which post-cranial bones are available to com- pare. The relative completeness of the holotype of O. howarthi permits us to make some additional osteological comparisons beyond those mentioned in the generic diagnosis. The fragments of cranium preserve evidence of the drepanidine affinities of Orthiospiza: the interorbital septurn has widely separated walls enclosing a network of bony trabeculae; the posterolateral margin of the cranial fenestra is straight; the anterior wall of the cranial cavity between the cranial fenestrae is broad and flat, not blade-like. Together these conditions define a synapomorphous character that occurs only in the Carduelinae, including the Drepanidini (Zusi 1978). The palatine process of the premaxilla is fused to the prepalatine bar in the configuration of a lateral flange, another derived character that is present in cardueline finches, but also appears in other, unrelated passerines (Book 1960). Orthiospiza has many presumably primitive characters in common with the other finch-billed drepanidine genera. The palatines have a very broad blade, short transpalatine processes and prepalatine bars, and have the walls of the palatine hasp wide-set to embrace the thick basisphenoid rostrum. The quadrate has the typical stocky form found in finch-billed drepanidines, with a thick body and a rounded medial condyle set on a short neck. The large narial opening in 50 ORNITHOLOGICAL MONOGRAPHS NO. 46 NEW HAWAIIAN BIRDS 51 A B C D FIG. 21. Selected postcranial bones of the holotype ofOrthiospiza howarthi, new genus and species (BPBM 160712). A, pelvis in dorsal view; B. left humerus in anconal view; C, right ulna in ventral view; D. right carpometacarpus in ventral view. Scale = I cm. Orthiospiza retains the subcircular shape that is typical of heavy-billed finches, rather than being elongated anteroposteriorly as is true of the relatively weak- billed Psittirostra and Melarnprosops, and of all thin-billed drepanindines except Hernignathus wilsoni and H. lucidus. The broad pelvis is also characteristic of the finch-billed as opposed to the thin-billed drepanidines. Autapomorphous characters that distinguish Orthiospiza from other drepani- dines are the greatly enlarged but not elongated narial openings, the absence of a medial trough on the maxilla, and the straight posteroventral edge of the maxilla. Orthiospiza shares at least one character with each of several other finch-billed genera, although none of these stands out as convincing evidence of relationship: the reduced size of the cranial fenestra is shared with Chloridops, the weak lateral nasal bars and pointed bill are shared with Rhodacanthis, and the weak ventral ridges that extend nearly to the tip of the maxilla resemble Loxioides and Telespiza. Two of the three known specimens of Orthiospiza are juveniles. These were not found in owl pellet deposits but appear to be remains of birds that entered FIG. 20. Mandibles ofOrthiospiza howarthL new genus and species, and Rhodacanthisfiaviceps. A, dorsal, E, lateral views of O. howarthL immature (USNM 372839); B, dorsal, D, lateral views of O. howarthœ holotype (BPBM 160712); C, dorsal, F, lateral views of R. fiavtceps, female {AMNH 453644). Scale = I cm. 52 ORNITHOLOGICAL MONOGRAPHS NO. 46 the caves unaided. A high proportion of juveniles dying in caves might be a reflection of species-specific behaviors such as cavity nesting or exploratory for- aging by young birds. Orthiospiza is one of relatively few avian fossil taxa that may have been re- stricted to montane habitats in the Hawaiian Islands. All of the specimens originate from three lava tubes located at relatively high elevations (1,145, 1,463, and 1,830 m). No specimens have yet been found in Puu Naio Cave (305 m), the most productive site for fossil passefines on Maui so far. It is a striking circumstance that the only two passefine skeletons preserved in the Upper Kipahulu Valley caves are of an extinct thrush and O. howarthi, while the abundant native species that occur in the vicinity now are not represented in the cave. Xestospiza, new genus Type species: Xestospiza fastigialis, new species. Included species: Xestospiza fastigialis, new species, Xestospiza conica, new species. Distribution: Kauai, Oahu, Molokai, and Maui. Etymology: Greek, xestos, planed, scraped, plus spiza, a finch; from the shape of the maxilla, which gives the appearance of having been planed to a straight profile, and in one species has a distinctly flattened dorsal surface. The gender is feminine. Diagnosis: Members of this genus are set apart from the other finch-billed drepanidines by their cone-shaped rather than arched bills. The dorsal profile of the maxilla is straight or nearly straight as opposed to distinctly curved as in all other finch-billed drepanidines. The bill is relatively long and narrow and the nafial openings are slightly elongated anteroposteriorly compared to the drepan- idine finches with heavier, arched bills (Telespiza, Rhodacanthis, Chloridops, and Orthiospiza). This genus most closely resembles Melamprosops, but differs in having a sturdier dorsal nasal bar, the maxilla deeper below the nafial openings, a shallower roedial groove and more pronounced ventral ridges, and a less deeply V-shaped postero- ventral contour of the maxilla, in addition to having a conical bill. Remarks: Because one of the new species incIuded in Xestospiza is known only from a pair of maxillae, we have limited the generic diagnosis to characters of that element. The conical rather than arched bill of this finch-like genus stretches the mor- phological breadth of the drepanidine radiation in a new direction. Finches with conical bills were widespread in the prehuman avifauna of the archipelago, as they occur on every island with a significant passefine fossil record. Although not necessarily each other's closest relatives, Xestospiza, Melampro- sops, and Psittirostra represent a similar grade of evolution within the drepanidine radiation in that each combines a finch-like bill shape with a few characters that are derived relative to the other finch-billed taxa. The morphological distance between Xestospiza and Melamprosops is not great, so that we were at first inclined to include the new species in the latter, to avoid introducing a new genus. We reconsidered when it proved that the only characters shared by the three species are ones that we would interpret as primitive. NEW HAWAIIAN BIRDS 53 TABLE 9 MEASUREMENTS (MM) OF 'rile MAXILLA IN Xestospiza, Melamprosops AND Psittirostra. H • HOLOTYPE Melamprosops Psittirostra Xestospiza conica Xestospiza fastigialis phaeosoma psittacea Dorsal length 16.9 H, 17.4 12.9 + H, 13.8, 14.6 15.1 16.4 Ventral length 10.5 H, 11.2 8.3, 8.4, 8.7 8.1 8.0 Length from lateral comer of nasofrontal hinge 16.0 H 13.3, 14.7, 14.7 14.9 16.2 Length from jugal articulation 12.9 H, 14.5 10.4, 10.7 H, 11.0 12.3 11.1, 11.9 Length from anterior narial opening 9.7 H, 9.9 7.4 H, 7.5, 7.9, 8.1 7.8 7.9 Maximum width 7.6 H 6.9, 7.2 H, 7.6, 7.7 7.2, 8.0 7.7, 8.1 Width of nasofrontal hinge -- 6.1 H, 6.8, 7.2 6.3, 7.0 8.2, 8.6 Length of narial opening 4.4 H 4.0, 4.1, 4.2 H, 4.3 4.3, 4.9 5.2, 5.3 Height of narial opening 3.5 H 2.9 H, 2.9, 3.1, 3.2 3.1, 3.3 3.6, 3.8 Height through lateral nasal bar 6.4 H 6.0 H, 6.1, 6.5 6.9, 7.6 7.4, 8.2 Minimum width of dorsal nasal bar 1.6 H, 1.8 1.3, 1.4, 1.4 1.1, 1.1 1.4, 1.6 Xestospiza conica, new species (Fig. 22G-J) "cf. Psittirostra, cone-billed finch" Olson and James, 1982b:40. "Psittirostra (subgenus incertae sedis), Cone-billed finch" Olson and James, 1982b:45; 1984:771. Holotype: Maxilla lacking the right lateral nasal bar, with a number of small holes worn through the premaxillary, USNM 254881 (Fig. 22G-I). Collected 15 August 1976 by Storrs L. Olson and Robin Rice. Type locality: Site K-3, Makewehi dunes, Kauai, Hawaiian Islands. Distribution: Kauai: known only from the type locality. Measurements ofholotype: See Table 9. Etymology: Latinized from Greek, konikos, conelike; from the shape of the maxilla. Paratype: Maxilla lacking lateral nasal bars, with the dorsal nasal bar broken and repaired, USNM 254951 (Fig. 22J). Measurements ofparatype: See Table 9. Diagnosis: The elongated maxilla of this finch is identified with Xestospiza by its nearly straight dorsal profile. It differs from that of X. fastigialis in having shorter ventral ridges and a much deeper medial trough on the ventral surface. It also lacks the flattened dorsosagittal surface and the rugose ridges on the dorsal nasal bar that distinguish X. fastigialis. The specimens ofX. conica are larger than maxillae of X. fastigialis (Table 9). Remarks: The holotype and paratype are the only specimens yet identified of X. conica. Our assignment of this species to the genus Xestospiza is tentative pending the discovery of a mandible. The owl pellet remains from Kauai also include two articular ends of mandibles representing two different species of finch- billed birds, one of which may well be X. conica, but the material is too fragmentary to determine which. 54 ORNITHOLOGICAL MONOGRAPHS NO. 46 NEW HAWAIIAN BIRDS 55 Xestospi•a fastigialis, new species (Figs. 22D-F, 23B, C, E, F, 24) "cf. Psittirostra, ridge-billed finch" Olson and James, 1982b:40. "Psittirostra (subgenus incertae sedis) Ridge-billed finch" Olson and James, 1982b:45; 1984:771. "Ridge-billed Finch" James, 1987:225. Holotype: Incomplete skeleton, USNM 445821. Collected 4 April 1988 by Storm L. Olson, Pauline Fiene-Severns, R. Michael Severns, and Thomas W. Stafford, Jr. The specimen consists of the maxilla (Fig. 24D-F), the mandible lacking a portion of the left ramus (Fig. 24A, C), a substantial fragment of the cranium (Fig. 24B), a fragment of the sternum preserving the manubrial spine and coracoidal sulci, part of the synsacrum, ten vertebrae, the dorsal half of both coracoids, the sternal end of the left coracold, the anterior half of the right scapula, the left scapula, the distal half of the right humerus, the left humerus (Fig. 24G), both ulnae (Fig. 24H), the proximal end of the right carpometacarpus, the left carpometacarpus (Fig. 24I), the left femur (Fig. 24J), the right tibiotarsus (Fig. 24K), the proximal end of the left tibiotarsus, the right tarsometatarsus (Fig. 24L), and a piece of the shaft of the left tarsometatarsus. Type locality: Crystal Cave (183 m), Manawainui Gulch, Maui, Hawaiian Is- lands. Distribution: Oahu: Barbers Point and Ulupau Head. Molokai: Moomomi dunes and Ilio Point. Maui: Puu Naio Cave, Lua Lepo, and Crystal Cave. Etymology: An adjective based on Latin, fastigium, a slope up or down to a point, a gable; from the shape formed by the converging ridges that ascend along the dorsal nasal bar in this species. Measurements (ram) of holotype: Length measurements, with the comparable measurement from Telespiza ultima (female, MVZ 124729) in parentheses: basi- temporal plate plus the occipital condyle, 4.8 (4.9); humerus, 17.4 (16.4); ulna, 17.5 (19.3); carpometacarpus, 9.9 (11.3); femur, 17.7 (16.9); tibiotarsus without the cnemial crest, 29.0 (28.8); tarsometatarsus, 21.3 (21.7). For additional mea- surements of holotype, see Tables 9 and 10. Paratypes: Oahu, Barbers Point: complete maxilla, USNM 322483; three slight- ly damaged mandibles, USNM 445799 (Fig. 23B, E), BBM-X 155522, and BBM-X 155728. Molokai, Ilio Point: ' Complete maxilla, BBM-X 178142. Maui, Crystal Cave, Manawainui Gulch: Partial skeleton, USNM 394085, consisting of the mandible lacking the posterior part of the left ramus (Fig. 23C, F), the basitemporal plate, a fragment of the sternum, the synsacrum, partial right coracoid, partial left scapula, distal end of left ulna, left carpometacarpus, two pelvic fragments including the acetabula, both femora and tibiotarsi, all somewhat damaged, and the left tarsometatarsus lacking the proximal end. Puu Naio Cave: Fxo. 22. Maxillae of Melamprosops and Xestospiza, new genus. A, lateral, B, dorsal and C, ventral views of Melarnprosops phaeosorna (ANMH 810456); D, lateral, E, dorsal and, F, ventral views of X. fastigialis, new species (USNM 445800; lateral view is of the left side printed in reverse); G, lateral, H, dorsal and,/, ventral views ofX. conica, new species, holotype (USNM 254881; lateral view is of the left side printed in reverse); J, dorsal view ofX. conica (USNM 254951). Scale = 1 cm. 56 ORNITHOLOGICAL MONOGRAPHS NO. 46 I.l.I Oh-, NEW HAWAIIAN BIRDS 57 TABLE 10 MEASUREMENTS (MM) OF THE •/La. NDIB• IN Xestospiza, Melamprosops AND Psittirostra Melamprosops Psittirostra Xestospiza f stigialis phaeosoma psittacea Total length Length of tomial crest Symphysis length Greatest width of symphysis Greatest width of mandible Symphysis height Length of lateral cotyla Width of articular end with medial process Height at lateral cotyla Height at angle of mandible Ramus length (middle part) 20.8 + H, 21.2, 21.6, 22.8 23.3, 24.1 23.8, 24.0 10.6, 11.1, 11.8 10.8, 11.0 8.7, 9.1 6.8 H, 7.6, 7.7, 8.4 7.1 5.2, 5.9 6.7, 6.8, 6.9, 7.1 H 7.5 7.4, 7.8 13.5 15.8 14.6, 14.8 2.5, 2.8 H, 2.8, 2.8 2.8 3.1, 3.6 2.1, 2.2, 2.2 2.6, 3.0 2.9, 3.1 3.7, 3.9 3.8, 4.7 4.2, 4.4 1.5, 1.5, 1.6 a, 1.6 1.5, 2.1 1.8, 1.9 3.7, 3.8 a, 3.8, 3.9 3.4, 4.0 4.6, 4.7 9.2 H, 9.2, 9.4, 9.9 11.4, 12.0 12.8, 13.4 complete maxilla with the frontal attached, USNM 445800 (Fig. 22D-F); slightly damaged mandible, USNM 445805; right quadrate lacking part of the orbital process, USNM 445801; left quadrate, USNM 445802. Measurements (ram) ofparatypes: Associated skeleton, USNM 394085: Cra- nium: length of basitemporal plate plus the occipital condyle, 4.8. Mandible: length, 21.1 (about 1.6 mm missing from the tip); height of symphysis, 2.9; width of articular end with medial process, 3.9; height at lateral cotyla, 1.8; height at the angle of the mandible, 4.0; length of ramus (middle part), 9.4. Carpometa- carpus: length, 9.7. Synsacrum: ventral length, 13.5. Femur: length, 17.4. Tibio- tarsus: length without cnemial crest, 28.8. Tarsometatarsus: estimated length, 20.2. For additional measurements of paratypes, see Tables 9 and 10. Diagnosis: A finch with a cone-shaped bill, distinguished from other drepani- dines by the two converging, rugose ridges that ascend along the dorsal nasal bar of the maxilla. Between these ridges, the dorsal surface of the nasal bar is fiat in cross-section, whereas this surface has a convex cross-section in other drepanidines except the long, thin-billed Aidemedia lutetiae, new genus and species. The ventral surface of the maxilla has a shallow medial trough and ventral ridges that are moderately developed posteriorly but become indistinct about three-quarters of the distance to the tip of the bill. This is in contrast to X. conica, which has a distinctly deeper medial trough and shorter ventral ridges. The mandible lacks retroarticular processes and in this and other respects is typically finch-like, with a thick-walled symphysis and a deep ramus, particularly at the angle of the mandible. However, the symphysis is relatively long and narrow and the articular end of the mandible is relatively small compared to other finch- billed drepanidines. The small lateral cotylae are slightly elevated and angled posteromedially. The middle part of the ramus is relatively short compared to Loxioides, Psittirostra, and Melamprosops, and in this respect is closer to the condition in the heavier-billed finches. The quadrate has the form typical of finch-billed drepanidines, with a stout body and a rounded medial condyle. In the holotype, the walls of the interorbital septurn are fused to form a single 58 ORNITHOLOGICAL MONOGRAPHS NO. 46 A D B C G H I d - L K FIG. 24. Selected elements of the holotype of Xestospizafastigialis, new genus and species (USNM 445821). A, dorsal view of mandible; B, ventral view of basicranium; C, lateral view of mandible; D, lateral, E, ventral and F, dorsal views of the maxilla; G, left humerus in anconal view; H, left ulna in ventral view; L left carpometacarpus in ventral view; J. left femur in posterior view; K, right tibiotarsus in anterior view;, L, right tarsometatarsus in posterior view. Scale = 1 cm. sheet of bone, as opposed to being entirely separated by an intervening network of trabeculae as they are in finch-billed drepanidines with heavier beaks such as the similar-sized T. ultima. The septurn is thus relatively weak and may even have been fenestrated in the missing anterior portion, as it sometimes is in NEW HAWAIIAN BIRDS 59 Melamprosops phaeosoma, a bird with similar development of the interorbital septum. Remarks: The body size is in the range of Telespiza ultima, although the distal wing elements may be somewhat reduced by comparison (see measurements of holotype). In general bill shape, Y(. fastigialis has an icterid-like aspect that is particularly reminiscent of cowbirds (Molothrus). Y(. fastigialis differs from Molothrus and other icterids in lacking a distinctive flange on the tomium of the mandible. Although the size of the skull in Y(. fastigialis is roughly the same as in M. ater, the interorbital septum is much less extensively fenestrated, with the optic foramen not connected anteriorly to an interorbital fenestra as in M. ater. Furthermore, the portion of the cranial fenestra that is preserved in the holotype shows a broad, flat posteroventral surface, as opposed to the blade-like, mounded surface seen in M. ater. These traits of the cranial fenestra and interorbital septum argue that Y(. fastigialis is a cardueline, not an icterid (Zusi 1978). Convergence of drepanidine bill morphology toward that oficterids is also exhibited by the new genus of thin- billed gaping birds, Aidemedia, and by Loxops sagittirostris. Fossils of Y(. fastigialis occur commonly on the three adjacent islands of Oahu, Molokai, and Maui, making this the most widely-distributed of the new fossil species. Possibly its former distribution was similar to that ofPsittirostra psittacea, Himatione sanguinea, and Vestiaria coccinea, which occurred historically on all six of the larger islands without exhibiting geographic differentiation in phenotypic characters. The disappearance of Y(. fastigialis throughout its range serves as a reminder that extinction affected broadly distributed species along with the more restricted ones. Genus Incertae Sedis, Additional Oahu Finch "cf. Psittirostra, additional Oahu Finch" Olson and James, 1982b:40. "Psittirostra (subgenus incertae sedis) additional Oahu finch" Olson and James, 1982b:45. Material: Right articular end of a mandible, USNM 255351. Distribution: Oahu: Barbers Point. Measurements (ram): Length of lateral cotyla, 3.5; length of lateral cotyla plus retroarticular process, 4.2; height of mandible at lateral cotyla, 2.0. Remarks: In this fragment of a sturdy finch-like mandible, the retroarticular process has a most unusual form, consisting of a dorsally-projecting knob-like process rather than the expected posteriorly-projecting blade-like process. Assum- ing we have correctly attributed the bone to the Drepanidini, there is no doubt that it represents an undescribed species. We were able to rule out Dysmorodre- panis munroi after we removed the skull and mandible from the unique holotype of this species (James et al. 1989), and we had previously eliminated all other contenders. Formal diagnosis of the species must be deferred until better speci- mens of it are found. Genus Hemignathus Lichtenstein, 1839 In certain recent sources (Berger 1981; Pureerican Ornithologists' Union 1983; Pratt et al. 1987), the amakihis (Loxops stejnegerL . viren& L. parms, and L. 60 ORNITHOLOGICAL MONOGRAPHS NO. 46 sagittirostris) have been merged with the genus Hemignathus, following Pratt (1979). We disagree with this and concur with earlier writers (e.g., Wilson and Evans 1890-1899; Perkins 1903; Areadon 1950), who included only the akialoas and "heterobills" in Hemignathus. These species are distinguished from other drepanidines by their very long and thin sickle-shaped premaxillae, with the nasals dramatically constricted anteriorly, and by possessing a greatly enlarged roedial groove on the ventral maxilla (Olson and James 1988). The roedial groove may function as a conduit for blood vessels and nerves, and its great size in Hemi- gnatbus may indicate that the tip of the bill in these probing birds experiences relatively rapid growth (Richard L. Zusi, pets. comm.). The amakihis are further distinguished from Hemignathus in lacking the plantaris muscle (Raikow 1976, 1977). The species we include in Hemignathus are divisible into two distinct groups; the "heterobills" (H. lucidus and H. wilsoni), which have the mandibular rostrum only about half as long as the maxillary rostrum, and the akialoas (H. stejnegeri, H. lichtensteini, H. lanaiensis, and H. obscurus),. which have the mandibular rostrum nearly as long as the maxillary rostrum. The morphological distance between these groups is great enough that they arguably belong in separate genera. In an earlier paper (Olson and James 1982b) we separated them as Hemignathus and Heterorhynchus, as had some previous authors, although we now realize that this nomenclatural treatment is erroneous because both generic names have the same type species. The following new species has a long mandibular symphysis. Hemlgna, thus upupirostris, new species (Fig. 25B-E) "Hoopoe-like sickle-bill" Olson and James, 1982b:41, 45; 1984:771; James, 1987:225. Holotype: Mandible lacking most of both articular ends, USNM 254171 (Fig. 25B, D). Collected 17 August 1976 by Storrs L. Olson, C. John Ralph, Carol P. Ralph, and John Luther. Type locality.' Site K-2, Makawehi dunes, Kauai, Hawaiian Islands. Distribution: Kauai: Makawehi dunes. Oahu: Barbers Point. Etymology.' "Hoopoe-billed," from Latin, upupa, the boopoe, plus rostrum, beak; in reference to the convergent similarity of the mandible to that of Upupa epops. Measurements (ram) of holotype: Length of tomial crest, 25.9+; symphysis length, 20.1 +; greatest width of symphysis, 4.3; symphysis height, 1.6; height at lateral cotyla, 1.4; height at angle of mandible, 2.0; ramus length (middle part), 13.2. • ß Paratypes; Oahu, Barbers Point: two fragments ofmandibular symphyses, USNM 255211 (Fig. 25C) and 255304 (Fig. 25E). Measurements ofparatypes: No meaningful measurements can be taken of the paratypes because of their fragmentary nature. Diagnosis: A bird with a very long, attenuated and decurved mandible, resem- bling that of Hemignathus s, tejnqgeri, H. lichtensteini, H. lanaiensis, and H. pro- cerus, as opposed to the heterobills, H. lucidus and H. wilsoni. This species is distinguished from all other drepanidines by having virtually lost the lingual trough in the mandibular symphysis, this .trough being extremely shallow and NEW HAWAIIAN BIRDS 61 A E F FIG. 25. Mandibles of Hemignathus stejnegeri (H. procerus auct.) and H. upupirostris, new species. A. lateral and F. dorsal views of H. stejnegeri (USNM 19094); B. lateral and D. dorsal views of H. upupirostris. holotype (USNM 254171: lateral view is of the left side printed in reverse); C. dorsal view ofproximal portion ofsymphysis ofH. upupirostris (USNM 255211); E. dorsal view ofsymphysis lacking tip of H. upupirostris (USNM 255304). Scale = I cm. limited in extent to the caudal three-eighths of the elongated symphysis. The anterior five-eighths ofthe dorsal surface of the mandible has a flat cross-section instead of being deeply excavated. The mandible also differs from that in a skeleton of H. stejnegeri n being slightly larger, with the coronoid processes di placed anteriorly. It is much longer than the mandibles of H. lucidus or H. wilsoni. The portion of the articular end 62 ORNITHOLOGICAL MONOGRAPHS NO. 46 preserved in the holotype does not exhibit the modifications for hammering that are apparent in H. wilsoni (Zusi 1987). Remarks: The holotype and two paratypes are the only specimens that we have identified as H. upupirostris. We have not succeeded in associating a maxilla with this species: either it is morphologically inseparable from H. stejnegeri or we have not collected it yet. It might be questioned whether the fossils we have described as H. upupirostris are not in fact from the extinct Oahu Akialoa, H. lichtensteini. This is not the case, however, because the only two existing specimens ofH. lichtensteini (ZMB 7918, holotype; ANSP 3360, previously misidentified as H. stejnegeri) have a pronounced lingual trough in the mandible (pers. observ.). We infer from the shape of the mandible that the tongue ofH. upupirostris was shorter than the long, tubular organ in H. obscurus or H. stejnegeri. Lacking a trough on the anterior part of the mandible, the bill probably could not have accommodated such a long tongue. Although no other drepanidine lacks a well-developed trough on the dorsal surface of the mandible, this condition does occur in the hoopoes, woodhoopoes, and scythebills (Upupidae, Phoeniculidae, and Dendrocolaptinae: Campylorham- phus), with which H. upupirostris shares a convergent bill morphology (Olson and James 1982b). Vangulifer, new genus Type species: Vangulifer mirandus, new species. Included species: Vangulifer mirandus, new species; Vangulifer neophasis, new species. Distribution: Maul: lava tubes on the southern slopes of Haleakala Volcano. Etymology: "Little shovel bearer," from Latin, vangula, diminutive of vanga, a shovel, and the suffix -ifer, bearer, in allusion to the peculiar, blunt, somewhat spatulare appearance of the bill. The gender is masculine. Diagnosis: Drepanidines with long and delicate, rather than finch-like bills, distinguished from all other passerines we have examined by the long and broad, fairly deep bill culminating in a very blunt, rounded tip. Among drepanidines, the combination of a long bill with a somewhat rounded tip occurs also in Ai- demedia, from which Vangulifer differs in having the bill shorter and broader, with a slight lateral constriction of the maxilla, and a much shorter etroarticular process of the mandible. Vangulifer differs further from the finch-billed drepanidines in having antero- posteriorly elongated narial openings, very weakly developed and posterolaterally confined lateral ridges on the ventral maxilla, and in having a delicate mandible with the symphysis long and thin-walled and the middle part of the ramus narrow and straight. The middle part of the ramus is not abruptly deflected ownward, as it is in the highly nectarivorous drepanidines in Perkins' (1903) Division 1 (Himatione, Vestiaria, Palmeria, Drepanis, and Ciridops). The retroarticular pro- cess is weakly developed as compared to those genera, and to Loxops and Aide- media. As in some finch-billed and most thin-billed drepanidines, the postero- ventral edge of the maxilla is V-shaped. The roedial groove on the ventral maxilla exhibits a peculiar development: The main groove that in other genera extends straight from the posterior edge to the tip, is shallow in Vangulifer and is splayed NEW HAWAIIAN BIRDS 63 anteriorly into numerous accessory grooves that extend outward toward the lateral edges of the bone. Remarks: Despite the reduced size of the articular end of the mandible in V. mirandus, Vangulifer is unlikely to belong with the species in Perkins' Division 1, which are set apart by their deflected mandibular rami, long and delicate retroarticular processes of the mandibles, and their more laterally constricted and pointed bills. The various morphologies that are accommodated within the genus Loxops are either more finch-like (e.g., L. coccineus) or more similar to the species of Division 1, than is that of Vangulifer. Vangulifer may be related to Aidemedia, which has a blunt bill tip and some evidence of weak splaying of the medial groove. Other possible relatives are Paroreomyza, which lacks a pronounced retroarticular process and has a very delicate, yet pointed bill, and the Kauai Creeper Oreomystis bairdi, in which the retroarticular process is shaped like that of V. neophasis. Because Vangulifer can never be observed in life, it is impossible to know exactly how the blunt bill was used in foraging. Although the most reasonable guess might be derived through analogy with living forms that exhibit closely similar morphology, we have so far failed to find a satisfactory living analog for Vangulifer. Its bill seems too long and weak to be designed for seed cracking; too deep and broad to be suited for probing in bark; too blunt for a nectarivore; and with its weak retroarticular processes, unlikely to have been used for forceful gaping. Thrushes have broad, weak bills, but these are more pointed than in Vangulifer. Although we have entertained the possibility that Vangulifer used its bill to snap up insects on the wing, todies (Todidae) and flycatchers (Tyrannidae) exhibit more dorsoventral compression of the bill than does Vangulifer (see ad- ditional discussion in the remarks under V. mirandus). A clue to the use of the bill may exist in the distinctive splaying of the medial groove on the ventral surface of the maxilla. This might function to distribute the blood supply and/or nerves over the entire ventral surface of the maxilla, or to its lateral edges, rather than directly to the tip. No associated complete or partial skeletons of Vangulifer have been found. In identifying the following two new species, we have therefore relied on the con- formiry of the jaws and on the distribution of each morphotype at the various cave localities to predict which maxillae probably belong with which mandibles. We identified two maxillae and seven partial mandibles from Puu Naio Cave, and one maxilla from Lua Impo, as V. mirandus (minimum 5 individuals). To V. neophasis, we assigned 3 maxillae and 2 mandibular symphyses from Puu Makua Cave, one mandible from Lua Lepo, and 4 maxillae and 12 partial man- dibles from Puu Naio Cave (minimum 9 individuals). Vangullfer tnirandus, new species (Figs. 26D-F, 27B, D) "two new thin-billed drepanidine species" (part), James et al., 1987:2353. Holotype: Maxilla lacking the left lateral nasal bar, USNM 445807 (Fig. 26D- F). Collected 23 March 1988 by H. F. James. Type locality: Puu Naio Cave (305 m), Maui, Hawaiian Islands. Distribution: Maui: Puu Naio Cave and Lua Lepo. Etymology: Latin, mirandus, wonderful, strange. 64 ORNITHOLOGICAL MONOGRAPHS NO. 46 A D FIG. 26. Maxillae of Vangulifer, new genus. A, lateral, B. dorsal and, C, ventral views of V. neophasis, new species, holotype (USNM 44706 I); D, lateral, E, dorsal and F, ventral views of V. mirandus, new species, holotype (USNM 445807). Scale = I cm. Measurements (ram) o. fholotype: Dorsal length, 18.4; ventral length, 12.4; length from lateral corner ofnasofrontal hinge, 18.9; length from jugal articulation, 15.7; length from anterior margin of narial opening, 11.5; maximum width, approxi- mately 7.4; length of narial opening, 5.4; height of narial opening, 3.1; height through lateral nasal bar, 5.8; minimum width of dorsal nasal bar, 1.0. Paratypes: Maxilla lacking the nasal bars, USNM 445806; slightly damaged mandible consisting of the symphysis and the right ramus, USNM 445808 (Fig. 27B, D). Measurements (ram) o. fparatypes: Maxilla: ventral length, 11.2; length from jugal articulation, 16.0; length from anterior margin ofnarial opening, 11.1; max- imum width, 7.2. Mandible: total length, 28.2; length oftomial crest, 14.6; sym- physis length, 9.8; greatest width ofsymphysis, 6.3 +; symphysis height, 2.4; length of lateral cotyla, 2.8; length of lateral cotyla plus retroarticular process, 3.0; height at lateral cotyla, 1.4; height at angle of mandible, 2.6; length of ramus (middle part), 12.1. Diagnosis: Compared to Vangul•fer neophasis, the maxilla is more decurved and not nearly as deep, the narial opening is much smaller, the entire ventral surface of the maxilla is much less excavated and is more distinctly pitted antero- medially, the splaying of the medial groove is more pronounced, and the tomial NEW HAWAIIAN BIRDS 65 D FIG. 27. Mandibles of Vangulifer, new genus. A, lateral and C, dorsal views of V. neophasis, new species (USNM 445813; lateral view is of the left side printed in reverse); B, lateral and D, dorsal views of V. rnirandus, new species (USNM 445808). Scale = I cm. crest is flared laterally toward the jugal articulations, as opposed to being straight. While the mandible is similar in overall size to that of V. neophasis, the articular end is smaller and the retroarticular process is much shorter. Remarks: In several characters, V. mirandus is more similar to aerial insecti- vores than is V. neophasis. The maxilla is not as deep, its ventral surface is nearly flat rather than domed, and the articular end of the mandible is reduced in size. However, a curved bill would not be expected in a bird that fed exclusively by catching insects on the wing. Vangulifer neophasis, new species (Figs. 26A-C, 27A, C) "two new thin-billed drepanidine species" (part), James et al., 1987:2353. Holotype: Slightly damaged maxilla, USNM 447061 (Fig. 26A-C). Collected 10 June 1988 by H. F. James. 66 ORNITHOLOGICAL MONOGRAPHS NO. 46 Type locality: Puu Naio Cave (305 m), Maul, Hawaiian Islands. Distribution: East Matfl: Puu Naio Cave, Lua Lepo, and Puu Makua Cave. Etymology: Greek, neos, new, plus phasis, appearance, look; from the novel appearance of the bill. The name is a feminine noun in apposition. Measurements (mm) ofholotype: Dorsal length, 19.6; ventral length, 10.7; length from lateral comer of nasofrontal hinge, 19.8; length from jugal articulation, 16.5; length from anterior rim of narial opening, 7.7; ventral width, 7.6; length of narial opening, 7.7; height of narial opening, 3.8; height through lateral nasal bar, 7.2; minimum width of dorsal nasal bar, 1.2. Paratypes: Puu Naio Cave: maxilla lacking the left nasal bar, USNM 445809; mandible lacking the posterior half of the right ramus, USNM 445813 (Fig. 27A, C). Puu Makua Cave: Three damaged maxillae, USNM 445810, 445811, and 445812. Measurements (mm) ofparatypes: Maxillae (in the order listed above): dorsal length, 20.8+, 20.6+, --, --; ventral length, 11.5, 11.8, --, --; length from lateral comer of nasofrontal hinge, 20.8, 20.7, --, --; length from jugal articulation, 17.3, 17.1, , ; length from anterior rim of narial opening, 10.9, 10.6, --, --; max- imum width, 7.5, 8.0, --, 7.3; length of narial opening, 7.7, 6.9, 7.7, 7.6; height of narial opening, 4.2, 3.7, 4.0, 3.7; height through lateral nasal bar, 7.8+, 7.7+, --, 8.0+; minimum width of dorsal nasal bar, 1.2, 1.2, 1.2, 1.0, Mandible (approximately 2 mm should be added to length measurements to compensate for the damaged tip): total length, 29.6+; length of tomial crest, 16.1 +; symphysis length, 9.8 +; greatest width of symphysis, 6.8; symphysis height, 2.6; width of articular end with roedial process, 4.4; height at lateral cotyla, 1.8; height at angle of mandible, 2.9; length of ramus (middle part), 11.8. Diagnosis: Compared to V. mirandus, the maxilla is straighter and deeper, the narial openings are larger, the entire ventral surface of the maxilla is much more excavated, and the tomial crests are not flared outward towards the jugal artic- ulations. The ventral surface of the maxilla is less distinctly pitted anteromedially, and the splaying of the roedial groove is less pronounced. The mandibular artic- ulation is relatively large and has a longer retroarticular process compared to V. mirandus. While the retroarticular process exhibits individual variation in depth, it is shorter and sturdier than in Loxops, Himatione, Vestiaria, Palmeria, Drepanis, or Ciridops. This is a medium-sized drepanidine, the mandibular ramus posterior to the tomial crest be'rag comparable in length to Palmeria dolei. Remarks: The differences between V. neophasis and V. mirandus are great enough that these species might have been described as separate genera. The shared characters that influenced our decision to unite them are the blunt bill, splayed roedial groove, and short retroarticular processes, combined with a del- icate, elongate bill. Aidemedia, new genus Type species: Aidemedia lutetiae, new species. Included species: Aidemedia lutetiae, new species; A. chascax, new species; A. zanclops, new species. Distribution: Oahu, Molokai, and Matfl. NEW HAWAIIAN BIRDS 67 Etymology: To Joan Aidem, resident of Molokai and pioneer collector of fossil birds in the Hawaiian Islands. The word has no relationship with the Greek root aidemon, meaning bashful or modest (cf. Aidemonia, Nectadniidae; Aidemosyne, Estrildidae). The unusual terminal orthography results from our inability to resist creating a palindrome. The name is to be regarded as feminine in gender. Diagnosis: Drepanidine birds with sturdy, straight or decurved, very elongate bills, with extremely long retroarticular processes on the mandibles. Within dre- panidines, the combination of an elongate bill and moderately to well-developed retroarticular processes also occurs in Hemignathus, Loxops, Himatione, Pal- meria, Vestiaria, and Drepanis. Aidemedia differs from these in having the bill narrower posteriorly; the maxilla is nearly uniform in breadth as opposed to being expanded posteriorly; and the dorsal nasal bar is also of uniform breadth, rather than broadening anteriorly as in the other genera. The retroarticular processes are longer and sturdier than in any of the aforementioned taxa except Loxops sagitti- rostris. Aidemedia is tinther distinguished from Hemignathus in having a much less attenuated maxilla with a shallower medial groove. Aidemedia differs further from Himatione, Palmeria, Vestiaria, and Drepanis in the sturdier construction of the bill, the relatively large articular end of the mandible, the fairly straight rather than abruptly deflected anterior portion of the mandibular ramus, and the less excavated ventral surface of the maxilla. Aidemedia differs further from Lox- ops in having the anterior edge of the narial opening not defined by a clear rim. Among drepanidines, Aidemedia is most similar in general bill morphology to Loxops sagittirostris, but differs in the characters mentioned above and also in the larger bill and less pronounced retroarticular notch on the mandible. The maxilla has the ventral ridges and lateral troughs weakly developed and confined posterolaterally, with a moderately excavated medial trough occupying most of the ventral surface. These conditions are typical of long-billed drepani- dines. The medial groove is shallow with some weakly defined accessory grooves radiating from it anteriorly in a configuration that resembles the distinctly splayed groove of Vangulifer mirandus. Remarks: Pronounced retroarticular processes on the mandible occur in birds that employ forceful gaping to obtain food, including a variety ofpasserine species. Beether (1951) describes the use of gaping by Sturnus vulgaris and various icterids to feed in such diverse substrates as flowers, fruit, grass, bark, and earth. It seems likely that Aidemedia fed similarly. Despite its general similarity of bill shape to L. sagittirostris, Aidemedia exhibits some characters that suggest its relationships may lie elsewhere within the dre- panidine radiation. For instance, the flattened dorsal nasal bar is shared with Xestospizafastigialis, and the blunt bill tip and weak splaying of the medial groove of the maxilla are reminiscent of Vangulifer. Aidemedia chascax, new species (Figs. 28H, I, J, 29B, 30B) "Icterid-like gaper, Oahu" Olson and James, 1982b:41, 45; 1984:771. Holotype: Mandible with slightly damaged symphysis, lacking the right articular end and part of the right ramus (middle part), BBM-X 155523 (Figs. 29B, 30B). Collected July or August 1977 by Aki Sinoto and others. Type locality: Site 50-Oa-B6-100b, Barbers Point, Oahu, Hawaiian Islands. 68 ORNITHOLOGICAL MONOGRAPHS NO 46 I} H FIG. 28. Maxillae of Aidemedia, new genus, and Loxops sagittirostris. A. dorsal, B, lateral and C, ventral views of L. sagittirostris (AMNH 453236); D, dorsal, F, lateral and G. ventral views of A. lutetiae, new species (BPBM 175622); E, lateral view ofA. lutetiae (USNM 445816); H, dorsal, I, lateral, and J, ventral views ofA. chascax, new species (BBM-X 155844). Scale = 1 cm. NEW HAWAIIAN BIRDS 69 TABLE 11 MEASUREMENTS (MM) OF THE MAXILLA AND MANDIBLE IN Aidemedia COMPARED TO Loxops sagittirostris. n = HOLOTYPE Aidemedia Aidemedia Loxops Aidemedia lutetiae chascax zanclops sagittirostris Maxilla: Ventral length 17.3, 17.8, 18.2 20.0 -- 12.4 Length from jugal articulation 19.8, 20.0, 21.7 -- -- 15.7 Length from anterior narial opening 14.2, 16.6, 17.4 20.0 -- 11.6 Minimum width of dorsal nasal bar 1.6, 1.6, 1.7, 1.7 1.6 -- 1.0 Mandible: Total length 34.6, 36.4 }• 43.6 }• 39.8 }•* 30.2 Length of tomial crest 17.1, 19.6 }• 22.6 }• 24.6 }• 15.0 Symphysis length 13.2, 15.2 }• 17.5 }• 20.0 }• 10.7 Greatest width of syraphysis 5.5, 5.8 }• 6.3 }• 5.6 }• 6.3 Greatest width of mandible 14.4 }• -- -- 12.7 Syraphysis height 2.6, 3.0 }• 2.8 }• 2.3 }• 2.0 Length of lateral cotyla 3.9, 4.0, 4.1 }• 4.6 }• 4.0 }• 3.2 Length of lateral cotyla with retroarticular process 7.6 }•, 7.6, 7.7 8.6 }• -- 6.0 Width of articular end with raedial process 4.6, 5.0, 5.1 }• 5.5 }• -- 3.8 Height at lateral cotyla 2.4, 2.7 }•, 2.9 2.5 }• 2.3 }• 2.0 Height at angle of mandible 3.1, 3.5 H, 3.5 3.1 H 2.7 H 2.7 Ramus length (middle part) 10.8 H, 11.7, 11.7 13.8 H 12.9 H 10.6 * Approximately 4 mm should be added to the mandible length ofA. zanclops to compensate for the missing retroarticular process. Distribution: Oahu: Barbers Point. It is as yet uncertain whether fragmentary Pleistocene fossils assigned to Aidemedia ("icterid-like gaper" James 1987:225) from Ulupau Head on Oahu also belong to this species. Etymology: Greek, chaskax, a gaper; from the adaptation of the bill for gaping. The name is a masculine noun in apposition. Measurements of holotype: See Table 11. Paratypes: Maxilla lacking posterior portion, BBM-X 155844 (Fig. 28H, I, J); two nearly intact mandibular symphyses, USNM 445814 and BBM-X 154918; fragment of left side of mandible, USNM 255564; tip of mandibular symphysis, USNM 255125. Measurements ofparatypes: See Table 11. Diagnosis: The very straight, sturdy bill resembles that of Aidemedia lutetiae, but is longer and has the dorsal surface of the maxilla less flattened. Also, the retroarticular process is deeper and is displaced ventrally, and the mandibular ramus (middle part) is not angled downward as strongly. The mandible is broader and less decurved than in A. zanclops. Remarks: See the following species. Aidemedia zanclops, new species (Figs. 29C, 30A) "Sickle-billed gaper, Oahu" Olson and James, 1982b:41. "Sickle-billed gaper" Olson and James, 1982b:45; James, 1987:225. Holotype: Mandible lacking the posterior end of the right ramus, with the roedial and retroarticular processes missing from the left articular end, BBM-X 155160 70 ORNITHOLOGICAL MONOGRAPHS NO. 46 NEW HAWAIIAN BIRDS 71 (Figs. 29C, 30A). Collected July or August 1977 by Aki Sinoto and others. The left posterior ramus had been separated from the rest of the bone probably while still in the fossil site, and was originally assigned catalog number BBM-X 155177 before it was recognized and re-attached. BBM-X 155160 is the correct catalog number for the entire specimen. Type locality: Site 50-Oa-B6-78, Barbers Point, Oahu, Hawaiian Islands. Distribution: Oahu: known so far only from the type locality, although frag- mentary fossils from Ulupau Head may represent he same species (James 1987). Etymology: Greek, zanclon, a sickle, plus ops, face; from the long, decurved mandible. The name is a masculine noun in apposition. Measurements ofholotype: See Table 11. Diagnosis: The mandible differs from other species of Aidemedia in being nar- rower and more gracile, and in having a decurved rather than straight symphysis, although the degree of curvature is slight compared to other sickle-billed drepan- idines (Drepanis, Vestiaria, and Hemignathus). In its elongation the mandible resembles A. chascax more than A. lutetiae. Remarks: With its narrow, decurved bill, this is certainly the most divergent of the three species of Aidemedia, yet in being elongated the bill is rather unex- pectedly more similar to the syrupattic species A. chascax than to the evidently allopatric A. lutetiae. Possibly the straight-billed (A. chascax) and curved-billed (A. zanclops) forms of Aidemedia from Oahu are males and females of a single, highly dimorphic species. Among passerines, a comparable extreme of sexual dimorphism in bill shape is reached only by the Huia, Heteralocha acutirostris (Callaeidae), of New Zealand, so that the possibility of this occurring in Aidemedia must be considered slight, particularly since no such dimorphism appears to be present in A. lutetiae of Molokai and Maul. Aidemedia lutetiae, new species (Figs. 28D-G, 29A, 30C) "Icterid-like gaper, Molokai" Olson and James, 1982b:41, 45; 1984:772. Holotype: Mandible with slight damage to the tip and mandibular foramina, BBM-X 147441 (Figs. 29A, 30C). Collected between September 1972 and January 1974 by Joan Aidem. Type locality: Site 10, Moomomi dunes, Molokai, Hawaiian Islands. Distribution: Molokai: Moomoral dunes. Maul: Puu Naio Cave, Lua Lepo, and Puu Makua Cave. Etymology: Latin, of Lutetia, the ancient Gallic capital of the Parisii, in allusion to the fact that members of this species must have spent their lives in gapery. Measurements ofholotype: See Table 11. Paratypes: Molokai, Moomomi dunes: two maxillae, both lacking lateral nasal bars, BBM-X 152622 and BPBM 175622 (Fig. 28D, F, G); a mandible lacking the posterior half of the left ramus, USNM 445818. Maul, Puu Naio Cave: Maxilla lacking right lateral nasal bar, USNM 445815; the posterior halfofa right mandibular ramus, USNM 445817. Lua Lepo: Maxilla lacking the left lateral nasal bar, USNM 445816. Measurements of paratypes: See Table 11. Diagnosis: This species is distinguished from A. chascax and A. zanclops by its shorter, very straight bill and by having the dorsal maxilla flattened. The fiat 72 ORNITHOLOGICAL MONOGRAPHS NO 46 D FIG. 30. Mandibles of Aidemedia. new genus. and Loxops sagittirostris in dorsal view..4, .4. zanclops, new species, holotyl•e (BBM-X 155160); B, .4. chascax, new species, holotyl•e (BBM-X 155523); C, .4. lutetiae, new species, holotype (BBM-X 147441); D, L. sagittirostris (AMNH 453236). Scale = I cm. surface extends anteriorly from the dorsal nasal bar until it becomes indistinct about mid-way to the tip. A. lutetiae is further distinguished from A. chascax by having the mandibular symphysis angled slightly downward relative to the middle part of the ramus. NEW HAWAIIAN BIRDS 73 Remarks: The bill of A. lutetiae resembles that of meadowlarks of the genus Sturnella (Icteridae), which also have long, straight bills with pronounced retroar- ticular processes on the mandible and a similarly flattened upper surface of the maxilla. We are confident that this similarity of bill shapes is correctly attributed to convergence rather than to genealogical relationship. Icterids possess a dis- tinctive flange on the mandibular tomium that Aidemedia lacks. Characters of Aidemedia that occur commonly in drepanidines but not in icterids are the relatively long mandibular symphysis, the lack of a distinct intercotylar tubercle on the mandible, and the excavated ventral surface of the maxilla. Genus Ciridops Newton, 1892 The sole previously described species in this genus, Ciridops anna, was known from only five museum specimens, three taken on the island of Hawaii, and two of unknown origin. None were originally preserved as skeletons, so we removed bones from a skin and a trunk preserved in alcohol (see Comparative Material Examined), from which we learned that the pelvic appendage in Ciridops is much stouter than in related taxa (the other members of Perkins' 0903) Division l, i.e., Himatione, Palmeria, Vestiaria, and Drepanis). While the tarsometatarsus in some of the finch-billed drepanidines is as stout as in Ciridops, the tibiotarsus and femur of Ciridops are stouter than in any other drepanidine. Associated with the robust hindlimb of Ciridops are dorsally expanded iliac shields of the pelvis. Similar but more extreme modifications of the pelvis and hindlimb occur in the passefine genera Orthonyx (Baird 1985), Bowdleria, and Mohua (Olson 1990a, b). The very short, superficially finch-like bill, along with a fancied resemblance in plumage to the cardueline genus Leucosticte, led Richards and Bock (1973: 125) to propose that Ciridops is a primitive drepanidine. We do not find support for this idea in osteology. The maxilla and mandible of Ciridops are not like cardueline finches but are shortened versions of the thin, weak structures found in the nectarivorous genera Himatione, Palmeria, Vestiaria, and Drepanis. If the evidence for a cardueline origin of drepanidines is correct, then Ciridops and its relatives share a highly derived rather than a primitive osteology within the radiation. Ciridops and the other genera in Perkins' Division 1 are osteologically quite similar to Loxops, from which their bills can be distinguished by a suite of minor characters. In Division 1, the anterior edge of the narial opening is not defined by a clear rim (true also of Loxops parvus). Further, the ventral surface of the maxilla is more excavated, the articular end of the mandible is small and weak, and the mandibular foramen is enlarged, compared to species of Loxops with bills of similar size. In addition to its stout pelvic appendage, Ciridops can be distinguished from Himatione, Palmeria, Vestiaria, and Drepanis by its much shorter bill, constricted dorsal nasal bar, upturned retroarticular process of the mandible, deep mandibular ramus (middle part), and enlarged mandibular foramen. We have identified fossils of Ciridops from Kauai, Oahu, and Molokai. Contrary to our previous assessment (Olson and James 1982b:42), the scant fossil material found so far on Molokai may not differ from Ciridops anna, so it is not discussed further here. 74 ORNITHOLOGICAL MONOGRAPHS NO 46 B FIG. 31. Maxillae of Ciridops in lateral view. A, Ciridops sp., Oahu (BBM-X 155727);B, C. tenax, new species, holotype (USNM 254913); C, C. anna (MCZ 10995). Scale = 1 cm. Ciridops tenax, new species (Figs. 3lB, 32B, 33D, E, 34C, D, 35A-C, G-I) "Ciridops sp., Kauai" Olson and James, 1982b:42, 45; 1984:771. Holotype: Maxilla lacking fight lateral nasal bar, USNM 254913 (Figs. 3lB, 32B). Collected 17 August 1976 by Storrs L. Olson, C. J. Ralph, Carol P. Ralph, and John Luther. Type locality: Site K-2, Makawehi dunes, Kauai, Hawaii. Distribution: Kauai: Makewehi dunes. Etymology: Latin, tenant, holding firmly; so named for the muscular leg and large, presumably grasping foot possessed by members of this genus. Measurements ofholotype: See Table 12. Paratypes: Associated bones of one individual, USNM 254985, consisting of four fragments of the maxilla, the mandibular symphysis, fight femur still in articulation with the acetabulum, the proximal end of the right tibiotarsus without the cnemial crest, the distal end of the right tarsometatarsus, and one pedal phalanx. The bones were cemented in a nodule of calcareous sand in the config- uration of a regurgitated owl pellet (Olson and James 1982b:fig. 10). When the matrix was removed, this same pellet also yielded bones of one individual each of Loxops stejnegeri and L. parvus. Maxilla lacking nasals, USNM 254607; three mandibular symphyses, USNM 254157 (Figs. 33E, 34D), 254158,254159; right mandibular ramus, USNM 254969 (Figs. 33D, 34C); left articular end of the mandible lacking medial process, USNM 254971; synsacrum with the left half of the pelvis and the fight iliac crest still fused, USNM 445819 (Fig. 35A); synsacrum, USNM 445820; three right femora, 254034, 254043, 254046, six left femora, USNM 254035, 254044 (Fig. 35C), NEW HAWAIIAN BIRDS 75 B D FIG. 32. Maxillae of Ciridops in dorsal (left) and ventral (right) views. A, C. anna (MCZ 10995); B, C. tenax, new species. holotype (USNM 254913); C, Ciridops sp., Oahu (BBM-X 155727); D, dorsal view of Ciridops sp., Oahu (USNM 255415). Scale = I cm. TABLE 12 MEASUREMENTS (MM) OF THE MAXILLA AND MANDIBLE IN Ciridops. H = HOLOTYPE Ciridops Ciridops tenax Ciridops sp., Oahu anna Maxilla: Dorsal length 11.4 H -- 13.5 Length from jugal articulation 9.3 •, 9.9 8.6, 8.6, 9.2 9.4 Length from anterior narial opening 5.5 I•, 6.4 5.3, 5.4, 5.4 6.1 Length of narial opening 4.5 • -- 5.0 Height of narial opening 2.3 •{ -- 2.6 Height through lateral nasal bar 5.2 •{ -- 6.0 Minimum width of dorsal nasal bar 0.6 •{ 0.5 0.7 Mandible: Symphysis length 5.8, 6.0, 6.2, 6.4 4.6, 4.7, 4.8, 5.1 5.9 Greatest width of symphysis 4.1 -- 4.5 Symphysis height 1.5 -- 1.8 76 ORNITHOLOGICAL MONOGRAPHS NO. 46 TABLE 13 MEASUREMENTS (MM) OF THE POSTCRANIAL SKELETON I Ciridops. MEAN, STANDARD EVIATION, AND RANGE ARE GIVEN FOR SAMPLE SIZES GREATER THAN 4; DATA ARE LISTED INDIVIDUALLY FOR SAMPLE SIZES OF 4 OR FEWER Ciridops tenax Ciridops p., Oahu Ciridops anna Synsacrum length Femur length Femur, proximal width Femur, mid-shaft depth Femur, distal width Tibiotarsus length Tibiotarsus, distal width Tarsometatarsus length Tarsometatarsus, proximal width Tarsometatarsus, mid-shaft width Tarsometatarsus, distal width 10.6, 10.7 -- -- 13.4 _+ 0.53 13.5 14.8' 12.5-14.5 n= 10 3.0 + 0.06 3.4 3.8* 2.8-3.0 n=9 1.1 _+ 0.04 1.3 1.3' 1.1-1.2 n= 10 3.2 _+ 0.11 3.6 4.2* 3.0-3.3 n = 10 26.2 -- 29.0 2.7 -- 3.3 19.9 _+ 0.42 18.5, 18.6, 19.9 21.6 19.2-20.5 n=8 3.0 -+ 0.13 3.0, 3.0, 3.2 3.4 2.8-3.2 n=7 1.1 -+ 0.07 1.2, 1.3, 1.3 1.4 1.0-1.3 n= 10 2.3 _+ 0.08 2.2, 2.3, 2.4 2.5 2.1-2.4 n= 10 * Femur measurements for Ciridops anna were taken from a fossil from Molokai. 254045 (Fig. 35B), 254240, 254047, 254965; left tibiotarsus, USNM 254062 (Fig. 35G); three right tarsometatarsi, USNM 254079, 254080, 254082; six left tar- sometatarsi, USNM 254078 (Fig. 35H), 254081, 254083, 254085, 254086 (Fig. 35I), 254167. Measurements (mm) ofparatypes: Partial skeleton, USNM 254985: Mandible: length oftomial crest, 8.4; length ofsymphysis, 6.0. Femur: length, 14.5; proximal width, 3.0; mid-shaft depth, 1.2; distal width, 3.3. Tarsometatarsus: mid-shaft width, 1.1; distal width, 2.1. For additional measurements of paratypes see Tables 12 and 13. Diagnosis: The maxilla is similar to that of Ciridops anna, as is the mandible, with the following exceptions: the retroarticular process is longer, resembling a more upturned and somewhat shortened version of the retroarticular process in Vestiaria coccinea, the middle part of the ramus is not as deep, and the mandibular foramen is less enlarged. The femur, tibiotarsus, and tarsometatarsus are stouter than in Himatione, Vestiaria, Palmeria, or Drepanis, but not as stout as in Ciridops anna. NEW HAWAIIAN BIRDS 77 B C D FIG. 33. Mandibles of Ciridops in dorsal view. A, Ciridops sp., Oahu (BBM-X 155172); B, Ciridops sp., Oahu (USNM 255292, part of retroarticular process missing); C. C. anna (MCZ 10995); D.C. tenax, new species (USNM 254969, part ofretroarticular process missing); E, C. tenax, USNM 254157. Scale = I cm. Remarks: In all of the characters mentioned in the diagnosis, C. tenax is in- termediate between its more derived relative, C. anna. and the other taxa in Perkins' (1903) Division 1. Thus C. tenax stands out as the only new drepanidine species that we could interpret as a possible "missing link" between two distinct morphotypes. Bones of the hindlimb ofC. tenax are consistently smaller than in the specimens examined of C. anna, suggesting that C. tenaxwas a somewhat smaller bird overall (Table 13). Ciridops sp., Oahu (Figs. 31A, 32C, D, 33A, B, 34A, B, 35D, E, J, K) "Ciridops sp., Oahu" Olson and James, 1982b:42, 45; 1984: 771. "Ciridops sp." James, 1987:225. Material: Maxilla lacking dorsal and left lateral nasal bars, BBM-X 155727 (Figs. 31A, 32C). Three damaged maxillae, USNM 255176, 255415 (Fig. 32D), 255179. Three mandibular symphyses, each with part of one ramus attached, USNM 255039, 255458, BBM-X 155172 (Fig. 33A). Two mandibular symphyses, USNM 255209, 255429. Left mandibular ramus without symphysis, USNM 255292 (Fig. 33B). Right femur, USNM 255124 (Fig. 35E). Left femur with 78 ORNITHOLOGICAL MONOGRAPHS NO 46 B C D FiG. 34. Mandibles of Ciridops in lateral view. A, Ciridops sp., Oahu (USNM 255292; image is of the left side printed in reverse); B, Ciridops sp., Oahu (BBM-X 155172; image is of the left side printed in reverse); C, C. tenax, new species (USNM 254969); D, C. tenax (USNM 254157); E, C. anna (MCZ 10995). Scale = I cm. damaged distal end, BBM-X 155669 (Fig. 35D). Right tarsometatarsus, USNM 255087 (Fig. 35K). Two left tarsometatarsi, USNM 255257, BBM-X 155689 (Fig. 35J). Distribution: Oahu, Barbers Point. Measurements: See Tables 12 and 13. Remarks: Fossils of Ciridops from Oahu, which are encountered fairly frequently in the Barbers Point deposits, resemble C. anna in having the long bones of the hindlimb very stout, but differ in having the bill and the long bones slightly shorter (Tables 12 and 13, Figs. 32-34 and 35). With such meager comparative material of C. anna available, the importance of these differences is unclear. The Oahu bird is perhaps a distinct species, but for the present we have refrained from naming it. DISCUSSION An updated conspectus of the fossil and historic distributions of resident pas- se fine birds in the Hawaiian Islands appears in Table 14. Numerous extralimital distribution records for extant birds are not discussed in the text, including several previously unpublished records for Maui from our 1988 field season. These are NEW HAWAIIAN BIRDS 79 LIJ 80 ORNITHOLOGICAL MONOGRAPHS NO. 46 TABLE 14 FOSSIL AND HISTORICAL DISTRIBUTION OF ENDEMIC PASSERINE BIRDS IN THE HAWAIIAN ARCHIPELAGO. F, FOSSIL RECORD (INCLUDES ARCHAEOLOGICAL CONTEXTS). H, HISTORIC RECORD. LY, LAYSAN; N, NIHOA; K, KAUAI; O, OAHU; Mo, MOLOKAI; LN, LANAI; MA, MAUI; H, HAWAII Ly N K O Mo Ln Ma H Family Corvidae Comus impluviatus Corvus viriosus Corvus aft. hawaiiensis Corvus hawaiiensis Family Sylviidae Acrocephalus familiaris Family Myiagridae Chasiempis sandwichensis Family Muscicapidae Myadestes palmeri Myadestes myadestina Myadestes lanaiensis Myadestes sp., cf. lanaiensis Myadestes obscurus Family Meliphagidae Moho braccatus Moho apicalis Moho bishopi Moho sp., Maui Moho nobilis Chaetoptila aft. angustipluma cf. Chaetoptila, narrow-billed sp. Chaetoptila angustipluma Family Fringillidae, Tribe Drepanidini Telespiza cantans Telespiza ultima Telespiza persecutrix Telespiza ypsilon Telespiza aft. ypsilon, Maui Loxioides bailleui Chloridops sp., Kauai Chloridops wahl Chloridops sp., Maui Chloridops kona Chloridops regiskongi Rhodacanthis aft. fiaviceps Rhodacanthis aft. palmeri Rhodacanthis fiaviceps Rhodacanthis palmeri Orthiospiza howarthi Xestospiza conica Xestospiza fastigialis Melamprosops phaeosoma Psittirostra psittacea Various unidentified finches Dysmorodrepanis munroi Pseudonestor xanthophrys Hemignathus lucidus Hemignathus wilsoni Hemignathus stejnegeri Hemignathus lichtensteini Hemignathus sp., cf. lanaiensis Hemignathus lanaiensis H H H H F F H F F F F F H FH FH H F F F F F F F F F F F F FH H F F FH F H F F F F F F F F F FH H H F H FH FH F H H H H H H H H NEW HAWAIIAN BIRDS 81 TABLE 14 CONTINUED Ly N K O Mo Ln Ma H Hemignathus obscurus Hemignathus upupirostris F F Oreomystis bairdi FH Paroreomyza maculata FH Paroreomyza montana FH H FH Parorecmyza fiammea H Vangulifer mirandus F Vangulifer neophasis F Aidemedia chascax F Aidemedia zanclops F Aidemedia lutetiac F F Loxops caeruleirostris H Loxops coccineus H H Loxops rnana Loxops parvus FH Loxops stejnegeri FH Loxops virens FH H H H Loxops sagittirostris Vestiaria coccinea H FH H H FH Drepanis funerea H F Drepanis pacifica Himatione sanguinea H FH H H FH Himatione sp., cf. sanguinea F F Palmeria dolei H FH Ciridops tenax F Ciridops sp., Oahu F Ciridops cf. anna F Ciridops anna H H H H H Moho sp. (a meliphagid), Hemignathus cf. lanaiensis (an akialoa), and Drepanis funerea (Black Mamo). Further documentation of these records will be published elsewhere. All of the passedfine fossils we have examined can be attributed to one of the five families that are known historically from the main islands (Corvidae, Myi- agridae, Muscicapidae, Meliphagidae, and Fringillidae), thus revealing no pre- viously unknown colonizations of the archipelago. This is in strong contrast to the non-passedfne fossils, which include representatives of 8 to 10 previously unknown extinct lineages (Olson and James 1991). The colonizing species that gave rise to these lineages would have included a petrel (Pterodroma), an ibis (Apteribis), a duck or shelduck (moa-nalos), up to three geese of unknown affinities (Geochen, supernumerary Oahu goose, very large Hawaii goose), a large crake (Porzana), an eagle (Haliaeetus), a harrier (Circus), and a stdfgld owl (Grallistrix) (Olson and James 1991). The fossil record extends the distribution of Corvus westward from the island of Hawaii to include Maul, Molokai, and Oahu. There is no reason to doubt that crows once occurred on all of the main islands. The two new species, C. impluviatus and C. viriosus, have been identified only from fossil sites in dry, lowland settings, whereas the fossils from higher elevation sites on Maui and Hawaii, while not positively identified, may not differ from the extant Hawaiian Crow, Corvus hawaiiensis. 82 ORNITHOLOGICAL MONOGRAPHS NO. 46 A much more complete picture of the breadth of adaptive radiation in the Drepaniclini has emerged through the fossil record. Extinction took its heaviest toll among the finch-billed taxa, which account for half of the new species de- scribed. Six other new species had diverse bill shapes that were probably mainly adapted for insectivory, including two meadowlark-like gapers, two sickle-bills, and two with broad, blunt bills. Only one of the new species belongs with the red-and-black, primarily nectafivorous species (Division 1 of Perkins [ 1903]), so that the predominance of nectarivory in the histofically-known radiation is evi- dently an artifact of differential extinction. Other extinctions were also non-random, removing all of the flightless and raptofial species everywhere except on the island of Hawaii. (We exclude Asio flarnrneus, which is not endemic and probably did not colonize the islands until after the arrival of humans [Olson and James 1982b]). Looking at passefines alone, we see that a wide distribution within the archipelago was no guarantee against extinction: 7 of the 16 new species (440/0) are known from more than one island. Forest passe fines that feed on nectar and insects seem to have been the best survivors. We have examined almost all skin, skeleton, and alcoholic specimens of Ha- waiian birds in Europe, North America, and Hawaii, and have not detected any overlooked specimens of the species we have described from fossils. Nevertheless, some of the extinct species described here may have survived in small numbers into the early part of the historic period (i.e., between the first western contact in 1778 and the first systematic ornithological collecting, beginning in the 1880's). Although it may be unrealistic to expect to be able to identify specific causes of extinction for individual species (Diamond 1984), the chronological data linking rapid disappearance of a major portion of the Hawaiian avifauna with prehistoric human settlement is convincing. Factors that probably played a role in avian extinctions are habitat destruction, particularly of lowland forests, predation by humans and introduced mammals, and possibly unidentified introduced diseases (Olson and James 1982b, 1984, 1991). In considering the passefines by them- selves, it seems unlikely that human predation was a primary cause of extinction for most of the small forest species, although it may have played a larger role in the disappearance of crows. Nor were the particular mammalian predators that were introduced by the Polynesians (dogs, pigs, and the Pacific rat [Rattus exu- lans]) likely to have decimated tree nesting species. It is possible that some fraction of the extinct passe fine species (for instance, those in the genus Telespiza) were vulnerable to prehistoric mammalian predation because of nesting on or near the ground. Barring the discovery of devastating prehistoric introduction of avian disease, habitat alteration remains the leading candidate for the primary cause of extinction of so many small passefine species. The historically known avifauna of the Hawaiian Islands consists of 40 (lump- ing) to 55 (splitting) endemic species (Olson and James 1991), whereas 35 fossil species have now been diagnosed. While these fossil discoveries have improved our appreciation for the diversity of the prehuman avifauna of the islands, it is worth emphasizing that many more extinct species await description. We have refrained from formally naming 8 additional fossil forms of drepanidines, many of which will doubtless also prove to be new species, and there are a number of very fragmentary fossils, including the previously cited "Additional Kauai Finch" NEW HAWAIIAN BIRDS 83 (Olson and James 1982b), that we have deliberately omitted from discussion. Treatment of the Hawaiian fossils of Meliphagidae, of which there appear to be at least two new species present, is also deferred pending further study. In Part I (Olson and James 1991), we mention an additional 11 fossil forms that are potentially new species (an ibis, 5 anatids, and 5 rails). We can be certain there are more extinct species of which we have not collected even the first fragment, considering that the prehuman avifaunas of the islands of Niihau, Lanai, and Kahoolawe are essentially unknown, and there are major gaps in the record from the island of Hawaii. Although further collecting and descriptive work are still needed, the intro- duction of these new taxa ought to enable Hawaiian avian paleontology to emerge from what has of necessity been a purely descriptive phase. More attention can be now be given to other stimulating lines of research, such as the chronology, causes, and ecological consequences of extinction, and patterns in evolution and biogeography. ACKNOWLEDGMENTS The following acknowledgments apply to both parts of this study. We reiterate our indebtedness to the legions who have so generously assisted us with this project, both in the museum and the field, many of whom have already been mentioned in our Prodromus (Olson and James 1982b). We cannot let any op- portunity pass to thank again those who have repeatedly provided assistance and support in the islands: Joan Aidere, Allen Allison, Carla H. Kishinami, C. J. Ralph, Carol P. Ralph, and Alan Ziegler. The great effort contributed by the Department of Zoology of the Bernice P. Bishop Museum, Honolulu, in curating and lending the fossil bird material acquired through that institution is profoundly appreciated. The greatest advances that have been made in our knowledge of Hawaiian fossil birds since the publication of our Prodromus have come from Maui. We owe a special debt of gratitude to R. Michael Severns for his indefatigable efforts in locating fossil deposits on that island and for many hard days of work assisting us in collecting. For access to the majority of fossil sites on Maui, and for many other considerations that have facilitated our excavations, we are greatly indebted to Pardee Erdman, owner of Ulupalakua Ranch, and to Ed O. Rice, ranch foreman. In 1982 we were assisted on Maui by Joan Aidem, Tonnie L. C. Casey, Laura Johnson, Angela K. Kepler, Cameron B. Kepler, Patricia Lester, Ronald Lester, Dan Saunders, and R. Michael Severns. During our excavations on Maui in 1984 we enjoyed the expertise of Paul S. Martin, Patrick C. McCoy, and David S. Steadman. At that time we were also assisted (at times overwhelmed) by a mul- titude of volunteers, gathered largely by Angela K. Kepler and Cameron B. Kepler. We hope that we have not overlooked any of their names: Joan Aidere, Christina Berman, Lili Berman, John Bose, Phil Bose, Janet Butzine, Colin Cameron, Doug Cameron, Mary Cooke, Sam Cooke, Carmelle Crivellone, Bill Eichenlaub, Mary Evanson, Denny Fen, Ann Fielding, Betsy H. Gagnil, Bob Hobdy, Heinz Hoben, Hugo Huntzinger, Glen James, Willie Lansford, George LeBouvier, Lloyd L. Loope, Jonathan Mann, Grigoria Marrero, Nancy Mawson, Julia Murphy, Patty Nakao, Bruce Perrin, Sandy Powers, Lisa Raymond, David Salemme, Les Skill- ings, Mele Skillings, R. Michael Severns, Linda Stumley, Howell Thomas, Connie 84 ORNITHOLOGICAL MONOGRAPHS NO. 46 Uzunboylu, Curtis Walseth, Evelyn Walseth, Callyn D. Yorke. For part of the time during our prolonged excavations on Maul in 1988 we benefitted materially from the assistance of Thomas W. Stafford, Jr., in obtaining stratigraphic and radiometric data. For additional assistance in excavating, screening, removing bags of sediment, and other favors that facilitated our 1988 field work on Maul we thank: Joan Aidem, Sheldon Berman, Fern P. Duvall, II, Babs Faye, Pauline Fiene-Severns, Renate Gassman-Duvall, Robert C. Fleischer, Lawrence Harper, Francis G. 'Howarth, Sabine Jessel, Mr. and Mrs. Merton Kekiwi, Carla H. Kish- inami, Patty Lester, Ron Lester, Arthur C. Medeiros, Jr., Stephen Peteira, Thane Pratt, Ricky Purdy, Mike Rose, Ken Schmitt, R. Michael Severns, Katherine C. Smith, Thomas Smith, Sharon Thodt, Laura Thompson, Chuck Whiteman, Charles Wyndham, Edie Wyndham, and Clement Young. Collection of fossils on National Park Service land in the Kipahulu Valley, Maui; was made possible by: Legado Hanky Eharis, Anne Made LaRosa, Terrance Lind, Lloyd L. Loope, Arthur C. Medeiros, Jr., Ron Nagata, and Donald Reeser. We are most grateful for various forms of assistance in the paleontological exploration of the following islands: Kauai: Dave S. Boynton, Pauline Fiene- Severns, Reginald Gage, Sandy Gage, Hailer H. Hammatt, Mike Kido, William K. Kikuchi, R. Michael Severns. Oahu: Diane C. Ddgot, Robert J. Hommon, Carla H. Kishinami, Terry Lopez, Col. Charles Robinson, Thomas W. Stafford, Jr., Alan Ziegler. Molokai: Joan Aidem, Henry Laws, George K. Linney, Lloyd L. Loope, Earl Neller, Gary Somers. Kahoolawe: Tom Hauptman, various staff of the United States Navy. Hawaii: Newell Bohnett, William Cuccaro, John Earle, Jon G. Giftin, Francis G. Howarth, Jim Lavoie, Kathleen Lavoie, Stephen Moun- tainspring, Monty Richards, Nelson Santos, J. Michael Scott, Fred D. Stone. The museum aspects of this study would not have been possible without the loan of many valuable and extremely rare specimens, including study skins of extinct species from which we removed bones using the method described by Olson et al. (1987). The curators, technicians, and others who assisted us are identified here with the museum acronyms (see Materials and Methods): Wesley E. Lanyon and Lester L. Short (AMNH): Auckland University Geology Depart- ment (AU--specimens made available through a loan to P. R. Millener); P. J. K. Burton, Graham Cowles, and Ian Galbraith (BMNH); Allen Allison, Carla H. Kishinami, Robert Pyle, and Alan Ziegler (BPBM); Canterbury Museum (CMC-- specimens made available through a loan to P. R. Millener); Marion Jenkinson (KU); Raymond A. Paynter, Jr. (MCZ); Victoria M. Dziadosz, Ned K. Johnson, and Barbara Stein (MVZ--NSF Grant BMS 7200102); Herbert Schifter (NHMW); Sandy Bartie and P. R. Millener (NMNZ); Pierce Brodkorb (PB); A. Karkhu and E. N. Kurochkin (PIN); Gerlof Mees (RMNH); Molly Benson and Hilary Potter (UMZC); J. Phillip Angle (USNM); B. Stephan (ZMB). At the Smithsonian, we thank Frederick V. Grady, who for many years has assisted us in the preparation and sorting of Hawaiian fossil material. Mark Florence has worked patiently and diligently with us to catalog our immense fossil collections. Other assistance with collection management of fossils has come from Jonathan Becker, Susan M. Folkerth, and a grant from the Women's Committee of the Smithsonian Institution. Monetary support for our research in the Hawaiian Islands has otherwise come almost entirely through various funds and programs of the Smithsonian Institution, in which connection we express our great appre- NEW HAWAIIAN BIRDS 85 ciation to David Challinor, Robert Hoffmann, and S. Dillon Ripley. Victor E. Krantz of the Smithsonian Institution's Department of Photographic Services has spent countless hours with us over the years photographing Hawaiian and other fossils, for which are deeply appreciative. All of the photographs in the present publication are his work. The drawings in Figures 1 and 2 are by Jaquin B. Schulz. For checking the new names and their etymologies we are most grateful to George Steyskal, Department of Entomology, National Museum of Natural His- tory, Smithsonian Institution. For reading the manuscript and making many valuable comments we are indebted to Jonathan Becker, Ned K. Johnson, Robert J. Raikow, David W. Steadman, and Alan C. Ziegler. SUMMARY Along with the non-passerine birds described in Part I (Olson and James 1991), abundant fossils of passerines were collected in the Hawaiian Islands over the past fifteen years. We describe 4 new genera and 16 new species in the families Corvidae and Fringillidae, and mention, but refrain from describing, up to 8 additional new species. We do not treat the fossil records of Hawaiian thrushes (Myadestes), flycatchers (Chasiempis), or honeyeaters (Meliphagidae) in this pa- per. Distributions of the fossil and historically-known passefines in the Hawaiian Archipelago are summarized in Table 14. The major collections of passefines are from Kauai, Oahu, Molokai, and Maui, with less significant collections, mainly from archaeological contexts, available from the island of Hawaii. The other islands have no relevant fossil record as yet. The different modes of deposition of passefine and non-passerine fossils are briefly discussed. An extinct genus ofornithophagous owl, Grallistrix, contributed importantly to the fossil record by concentrating passefine remains at its roosts. The passefine fossil material described here dates to the Holocene (< 10,000 years ago). Fossils that date to over 120,000 years ago are known from Late Pleistocene sediments at Ulupau Head on Oahu, but we have not made use of this material in describing new taxa. We describe two new species of crows, both larger than the extant Corvus hawaiiensis. C. irnpluviatus, a new species with a high, arched bill, is known only from Oahu, while C. viriosus, a new species with a long, straight bill, occurs on both Oahu and Molokai. Four new genera and 14 new species of Drepanidini (Fringillidae: Carduelinae) are introduced, 8 with finch-like bills and 6 with a variety of more derived bill forms. Fossils of Telespiza, a genus of finches restricted in historic times to Laysan and Nihoa, are widespread in the main islands with up to three species occurring on a single island. Telespiza persecutrix, new species, is from Kauai and Oahu. T. ypsilon, new species, is from Molokai and Maui; and a very small specimen from Maui may represent yet another new species. T. cantans (Laysan Finch) and T. ultima (Nihoa Finch) also occur as fossils in the main islands. Chloridops kona (Kona Finch) from the island of Hawaii is the only historically known member its genus, but fossils reveal a greater diversity and wider distri- bution for Chloridops. C. wahl new species, is a smaller form from Oahu and Maui. A second, even more diminutive, species may be indicated on Maui. Chlor- 86 ORNITHOLOGICAL MONOGRAPHS NO. 46 idops sp. from Kauai resembles C. wahi but may prove to be a distinct species. The impressive new species C. regiskongi, from Oahu, is distinctive for the large size of its bill. Orthiospiza howarthi, new genus and species, is a finch known so far only from relatively high elevations on Mr. Haleakala, Maui. Apparently a resident of wetter forests, O. howarthi had unusually large nafial openings for such a heavy-billed finch. The species of Xestospiza, new genus, had finch-like bills with a straight rather than arched profile to the maxilla. Xestospiza conica, new species, is known only from Kauai, whereas X. fastigialis, new species, has the widest distribution of any new fossil form, being known so far from Oahu, Molokai, and Maul. We postpone describing four finch-billed drepanidines either because the fossils available would be inadequate as types or because larger series of comparative skeletons are needed to evaluate variation. These include two possible new species of Rhodacanthis (koa finches), and two distinctive finches of unknown generic affinites, one each from Oahu and Maul. Among the drepanidines that were not finch-like, Hemignathus upupirostris, new species, from Oahu and Kauai, is a sickle-billed species with a mandibular symphysis resembling that of hoopoes (Upupidae). Vangulifer is an enigmatic new genus from Maui, containing the new species V. mirandus and V. neophasis. These had rather long and weak bills that were remarkably broad anteriorly. What these birds may have fed on is a mystery. Aidemedia is a new genus of drepanidines adapted for gaping, of which we describe three new species, A. lutetiae from Molokai, A. chascax from Oahu, and A. zanclops, also from Oahu. The latter had a sickle-shaped bill while the first two had straight bills like those of meadowlarks (Ictefidae: $turnella). Of the 5 genera in Perkin's (1903) Division 1, only Ciridops has a greater diversity in the fossil record. We name C. tenax as a new species from Kauai, and discuss, but defer naming, a form of Ciridops from Oahu. Fossils of Ciridops from Molokai may prove to be conspecific with C. anna, known historically from the island of Hawaii. The extinction of so many species of Hawaiian passefines is attributed mainly to prehistoric human-wrought changes in forest habitats. LITERATURE CITED AM•r•o•q, D. 1950. The Hawaiian honeycreepers (Aves, Drepaniidae). Bull. Amer. Mus. Nat. Hist. 95(4): 151-262. AMERICAN ORNITHOLO(}ISTS' UNION 1983. Check-list of North American Birds, 6th ed. American Ornithologists' Union [Washington, D.C.]. BAn•, R.F. 1985. Arian fossils from Quaternary deposits in 'Green Waterhole Cave', southeastern South Australia. Rec. Austral. Mus. 37:353-370. 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WInSON, S. B., AND A. H. EVANS. 1890-1899. Aves Hawaiienses: The Birds of the Sandwich Islands. R. H. Porter, London. [A facsimile reprint was issued in 1974 as part of the series "Natural Sciences in America" by Arno Press, New York.] Z•SWILER, V. 1965. Zur Kennails des Samen6ffnens trod der Struktur des h6rnernen Gaumens bei kfrnerfressenden Oscines. J. Ornithol. 106(1): 1-48. Zusi, R.L. 1978. The interorbital septum in cardueline finches. Bull. Brit. Ornithol. Club 98(1): 5-10. Zus•, R.L. 1987. A feeding adaptation of the jaw articulation in new world jays (Corvidae). Auk 104(4):665-680. ORNITHOLOGICAL MONOGRAPHS No. I. A Distributional Study of the Birds of British Honduras. Stephen M. Russell. 1964. (Out of print) No. 2. A Comparative Study of Some Social Communication Patterns in the Pelecaniformes. G. F. van Tets. 1965. $2.50. No. 3. The Birds of Kentucky. R. M. Mengel. 1965. $10.00. No. 4. Ev•uti•n•fS•meA•cticGu•s(Larus):anExPerimenta•Study•fIs•atingMechanisms•Nea•G.Smith. 1966. (Out of print) No. 5. A Comparative Life-history Study of Four Species of Woodpeckers• Louise de Kiriline Lawrence. 1967. (Out of print) No. 6. Adaptations for Locomotion and Feeding in the Anhinga and the Double-crested Cormorant. O. T. Owre. 1967. $3.00. No. 7. A Distributional Survey of the Birds of Honduras. B. L. Monroe, Jr. 1968. $7.00. No. 8. An Approach to the Study of Ecological Relationships among Grassland Birds. John A. Wiens. 1969. (Out of print) No. 9. Mating Systems, Sexual Dimorphism, and the Role of Male North American Passerine Birds in the Nesting Cycle. Jared Verner and Mary F. Willson. 1969. (Out of print) No. 10. The Behavior of Spotted Antbirds• E. O. Willis. 1972. $4.00. No. 11. Behavior, Mimetic Songs and Song Dialects, and Relationships of the Parasitic Indlgoblrds (Vidua) of Africa. R. B. Payne. 1973. $6.00. No. 12. 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