The Coleopterists Bulletin 78(1): 69–75. 2024.
Lanyualtica hsui Lee and Konstantinov, a New Genus and Species of 
Flea Beetles (Coleoptera: Chrysomelidae: Galerucinae: Alticini)  
from Lanyu Island, Taiwan
Chi-Feng Lee
Applied Zoology Division
Taiwan Agricultural Research Institute
189 Chung-Cheng Road, Taichung 413 
Wufeng, TAIWAN
chifeng@tari.gov.tw
 https://orcid.org/0000-0003-1996-0557
and
Alexander Konstantinov
Systematic Entomology Laboratory, USDA, ARS
c/o National Museum of Natural History, MRC 168
Washington, DC 20560, USA
alex.konstantinov@usda.gov
 https://orcid.org/0000-0001-6578-6735
ABSTRACT
Lanyualtica hsui Lee and Konstantinov, new genus and species, is described from Lanyu Island, Taiwan. It belongs to 
the Tegyrius genus group and is most similar to Parategyrius Kimoto and Gressitt. 
Keywords: flea beetles, Tegyrius, Chionanthus ramiflorus, Oleaceae
DOI.org/10.1649/0010-065X-78.1.69
Zoobank.org/urn:lsid:zoobank.org:pub:D6F6BE2D-3502-4539-83FC-133486D8059D
INTRODUCTION flavicornis (Baly). One previously unknown species 
belonging to this genus group was collected on 
Lanyu Island, a small, volcanic island off the Lanyu Island and is described herein as a new genus 
southeastern coast of Taiwan, is home to a number and new species.
of endemic plants and animals, including leaf bee-
tles. Among Chrysomelidae, three galerucines, 
Aulacophora kotoensis Chûjô, 1962 (Lee and MATERIAL AND METHODS
Beenen 2015; Reid et al. 2021), Lanolepta fulva 
Kimoto, 1991, and Monolepta ongi Lee and Staines, Terminology for morphological structures fol-
2010; two eumolpines, Rhyparida formosana lows Duckett and Daza (2004). To prepare draw-
Aslam, 1968 (a replacement name for Rhyparida ings of the genitalia, the abdomens of adults were 
basileptoides Chûjô, 1956) and Rhyparida kotoensis separated and boiled in a 10% KOH solution, 
Chûjô, 1956; and one cryptocephaline, Coenobius cleared in distilled water, then mounted on micro-
yosionis Chûjô, 1954, are endemic to Lanyu Island. scope slides in glycerin for observation. Drawings 
The Tegyrius genus group was defined by were made using a Leica M165 stereomicroscope. 
Prathapan and Konstantinov (2001) and is com- Microscope slides were illustrated using a Nikon 
posed of the following genera: Bikasha Maulik, ECLIPSE 50i microscope. Digital images for Figs. 
Dentilabra Medvedev, Lanka Maulik, Neorthana 1G and 2 were taken with Macropod Pro photomac-
Medvedev, Parategyrius Kimoto and Gressitt, rography system (Macroscopic Solutions, LLC, 
Philogeus Jacoby, Sahyadrialtica Prathapan and Tolland, CT, USA), processed with Zerene Stacker, 
Konstantinov, Sphaeraltica Ohno, Tegyrius Jacoby, version 1.04, and edited with Adobe Photoshop 
and Tribolia Chen. Six members of this genus group Elements 2020.
occur in Taiwan: Bikasha collaris (Baly), Lanka Specimens were examined from the following 
bicolor (Chûjô), Lanka fulva (Chûjô), Lanka minor collections: Natural History Museum, London, UK 
(Chûjô), Lanka nigra (Chûjô), and Sphaeraltica (BMNH), Taiwan Agricultural Research Institute, 
69
Downloaded From: https://bioone.org/journals/The-Coleopterists-Bulletin on 25 Mar 2024
Terms of Use: https://bioone.org/terms-of-use	Access provided by Smithsonian Libraries and Archives
70 THE COLEOPTERISTS BULLETIN 78(1), 2024 
Taichung, Taiwan (TARI), and National Museum of absent, without midfrontal sulcus, transverse- 
Natural History, Smithsonian Institution, Washington, oblique, wider than long, spindle-shaped; at a lower 
DC, USA (USNM). level than vertex and not entering interantennal 
space. Antennal calli and frontal ridge meeting at a 
R single point, forming a pit. Orbit narrow, supraor-ESULTS bital sulcus narrow and deep, continuing anteriad 
Tegyrius Genus Group as a broad, deep orbital sulcus. Subgenal suture well 
(sensu Prathapan and Konstantinov 2021) developed along base of mandible. Eyes diverging 
slightly basally, not emarginate near antennal sock-
Remarks. Members of this genus group possess et, vertical diameter 1.3–1.4 times transverse diam-
the following characters: a pair of subparallel ridges eter. Distance between eyes 3.2–3.4 times diameter 
present on the intercoxal part of the first abdominal of a socket, 1.0–1.1 times transverse diameter of 
ventrite (Fig. 1G), ridges fused anteriorly and open one eye. Diameter of antennal socket 2.0 times dis-
towards the posterior, presenting the appearance of tance between eye and adjacent socket. Distance 
an upturned “U”; the metatibia characteristically between antennal sockets 1.2–1.5 times diameter of 
curved in dorsal view, with its distal region curving a socket. Frontal ridge narrow, its base triangular, 
away from the long axis of the body, its dorsal sur- well-defined with raised lateral margin, forming a 
face rather flat, with lateral and mesal margins shallow groove between eye and frontal triangle. 
armed with bristles towards the apex; general hab- Frontal ridge surface rugose. Frontoclypeal suture 
itus oblong in dorsal view and convex in lateral with three pairs of long setae. Antenna filiform, 
view, resembling that of Longitarsus Latreille, reaching middle of elytron when laid over prono-
Psylliodes Latreille, or Aphthona Chevrolat; length tum. Labrum broader than long, with three pair of 
ranging from 1.5–3.5 mm; procoxal cavities open setiferous pores arranged in a transverse row. 
behind; penultimate maxillary palpomere thicker Maxilla with apical palpomere acute, longer and 
than preceding or apical palpomere; posterior mar- narrower than penultimate palpomere. Labial palpus 
gin of metaventrite bisinuate, raised on either side with three palpomeres, I shortest, II and III equal 
of middle forming a pair of protuberances distally; in length.
metatarsomere I articulated on a callosity, visible in Pronotum convex, 1.4–1.5 times wider than long; 
lateral view, on apex of metatibia, as in Longitarsus; lacking obvious antebasal impression, but slightly 
metatarsomere I forming an inverted L-shape in flattened basally; parallel-sided; anterolateral cal-
relation to remaining tarsomeres in dry specimens; losity slender, more than 4.0 times longer than wide, 
vaginal palpi elongate, joined mesally in proximal oblique in dorsal view, pore situated posterior and 
half, with elongate sclerotizations anteriorly and dorsal face of callosity and associated with an 
posteriorly and with long apical setae, generally obtuse denticle; posterolateral callosity slightly 
resembling those of Longitarsus or Aphthona; sper- protruding laterally, with laterally situated pore; 
matheca with a distinct receptacle, pump, and posterior margin bisinuate, forming an indistinct 
gland duct. median lobe; pronotal punctures coarse, more 
These characters undoubtedly place the beetles distinct than those on vertex; disc with reticulate 
in question into the Tegyrius genus group. However, microsculpture. Procoxal cavities open behind. 
they do not fit within any of the known generic con- Prosternal intercoxal process reaching same level 
cepts and are described herein as a new genus and as procoxae, widening, apical margin convex and 
new species. covered with long setae, disc with dense and coarse 
Lanyualtica Lee and Konstantinov, new genus punctures, each bearing one long seta; narrowest 
zoobank.org/urn:lsid:zoobank.org:act: width of prosternal intercoxal process 2.8 times 
7FE5E51E-F8EB-4440-8892-47F306E526B2 smaller than distance between anterior margin 
(Figs. 1–3) and coxal cavity; 0.5 times length of prosternum. 
Mesosternal intercoxal process parallel-sided, with 
Description. Small, elongate-ovate, convex in concave posterior margin, disc with extremely 
lateral view, length 2.73–3.27 mm, width 1.39–1.64 coarse punctures in one transverse row along pos-
mm. Color black, non-metallic, appendages brown terior margin; distance from anterior margin of 
(Figs. 1A–F). mesoventrite to end of mesoventral intercoxal pro-
Head hypognathous. Vertex and frons together in cess 1.5 times width of mesoventral intercoxal pro-
lateral view forming a convex line, slightly arched cess; width of mesoventral intercoxal process 
between eyes and antennal sockets. In frontal view 1.7–1.8 times minimum distance from anterior mar-
(Fig. 2A), vertex convex with scarce fine punctures gin of mesoventrite to coxal cavity.
and weak transverse wrinkles. Supraorbital pore Elytra as broad as pronotum basally, broadening 
situated dorsad of orbital sulcus, with a long seta. at basal one-third, without basal callus but with de-
Antennal calli poorly developed, supracallinal sulci pression at apical one-third, humeral callus well 
Downloaded From: https://bioone.org/journals/The-Coleopterists-Bulletin on 25 Mar 2024
Terms of Use: https://bioone.org/terms-of-use	Access provided by Smithsonian Libraries and Archives
 THE COLEOPTERISTS BULLETIN 78(1), 2024 71
Fig. 1. Lanyualtica hsui, new genus and species. A) Male, dorsal view, B) Ditto, ventral view, C) Ditto, lateral 
view, D) Female, dorsal view, E) Ditto, ventral view, F) Ditto, lateral view, G) Abdominal ventrites 1 and 2, H) Metatarsi, 
dorsal view.
Downloaded From: https://bioone.org/journals/The-Coleopterists-Bulletin on 25 Mar 2024
Terms of Use: https://bioone.org/terms-of-use	Access provided by Smithsonian Libraries and Archives
72 THE COLEOPTERISTS BULLETIN 78(1), 2024 
Fig. 2.  Lanyualtica hsui, new genus and species. A) Head, frontal view, B) Pronotum, dorsal view.
developed; disc with coarse punctures, slightly Metatarsomere III bilobed, asymmetrical (Fig. 1H), 
smaller than those on pronotum (Fig. 2B), arranged mesal lobe smaller than lateral lobe.
into 11 longitudinal rows; with much smaller punc- Intercoxal part of abdominal ventrite 1 raised, 
tures in between. Elytral epipleuron extending nine- with well-developed subparallel ridges (Fig. 1G). 
tenths of elytron, almost reaching apex, outwardly Aedeagus (Figs. 3C, D) elongate; rectum membra-
oblique with maximum width less than that of me- nous; endophallus with apical sclerite slender, re-
sofemur. Hind wings fully developed. Metaventrite curved inwards, followed by a pair of longitudinal 
with anterior margin raised, convex towards poste- rows of short setae, then a pair of longitudinal rows 
rior, forming paired tubercles distinctly raised above of short teeth, and with one elongate and laterally 
level of metacoxa. flattened basal sclerite. Spermatheca (Fig. 3G) with 
Pro- and mesotibiae convex dorsally, with no distinct pump, receptacle, and duct. Duct not coiled. 
apical spine. Metafemur robust with anterior margin Vaginal palpi (Fig. 3E) elongate, slightly fused me-
distinctly more convex than posterior margin. dially, with a few long apical setae. Abdominal 
Metatibia characteristically curved in dorsal view, sternite VIII (Fig. 3F) in females weakly sclerotized 
distal region curving away from the long axis of the but with well-sclerotized spiculum.
body, widening gradually towards distal end, preapi- Sexual Dimorphism. Posterior margin of last 
cally narrowing; convex dorsally in proximal one- abdominal ventrite in males bisinuate, with a medi-
third, flat medially and distally; distinctly margined an lobe. In females, posterior margin of last abdom-
laterally with row of bristles in distal one-fourth; inal ventrite widely rounded, with a shallow incision 
gradually broadening apically in lateral view; mesal at middle.
margin indistinct in proximal one-fourth, appearing Type Species. Lanyualtica hsui Lee and 
distally with row of acute bristles at distal one- Konstantinov, new species.
fourth, these bristles similar to those on lateral mar- Etymology. Lanyualtica is derived from two 
gin. Metatibial spur articulated in middle of tibial words, namely Lanyu, referring to Lanyu Island, the 
apex, acute. Metatarsomere I longer than half length type locality of the type species, and Altica Geoffroy, 
of metatibia, longer than length of metatarsomeres the type genus for the tribe. The gender is feminine.
II to IV combined; densely covered with long setae Diagnosis. Among genera in the Tegyrius group, 
ventrally. Metatarsomeres II and III subequal Lanyualtica is closest to Parategyrius by possess-
in length, asymmetrical, apex extending laterally. ing the following characters: metatibial spine acute; 
Downloaded From: https://bioone.org/journals/The-Coleopterists-Bulletin on 25 Mar 2024
Terms of Use: https://bioone.org/terms-of-use	Access provided by Smithsonian Libraries and Archives
 THE COLEOPTERISTS BULLETIN 78(1), 2024 73
Fig. 3. Lanyualtica hsui, new genus and species. A) Antenna, male, B) Antenna, female, C) Aedeagus, dorsal view, 
D) Ditto, lateral view, E) Vaginal palpi, F) Abdominal sternite VIII, female, G) Spermatheca.
Downloaded From: https://bioone.org/journals/The-Coleopterists-Bulletin on 25 Mar 2024
Terms of Use: https://bioone.org/terms-of-use	Access provided by Smithsonian Libraries and Archives
74 THE COLEOPTERISTS BULLETIN 78(1), 2024 
anterofrontal ridge not abruptly raised above cly- becoming broader, VIII to XI with same width; an-
peus, not forming right angle in lateral view; an- tennomere III shortest, III to VII each slightly longer 
tennal calli not depressed, at same level as adjoining than preceding antennomere, VII to X with same 
area of vertex; frontal ridge more-or-less separated length, XI longest, ratio of length of antennomeres 
from vertex by antennal calli; metatarsomere I half I to XI 1.0:0.6:0.6:0.6:0.8:0.9:0.9:0.9:1.0:0.9:1.3; 
the length of the metatibia; and antennal calli reach- ratio of length to width of antennomeres I to XI 2.5:
ing but not entering deeply into the interantennal 1.9:2.5:1.8:2.3:2.4:2.5:2.0:2.2:1.8:2.8. Aedeagus 
space. Lanyualtica can be differentiated from (Figs. 3C, D) elongate, 6.0 times longer than wide, 
Parategyrius by the head lacking supracallinal apically narrowed from apical one-ninth, apex nar-
sulci, pronotum without transverse antebasal im- rowly rounded, slightly narrowed at apical one-
pression and with coarser punctures, and elytral fourth, parallel sided from apical one-third to base, 
punctures arranged in regular rows, some of which strongly narrowed at basal one-third, slightly curved 
are strongly depressed. This last character together in lateral view; rectum membranous; endophallus 
with asymmetrical metatarsomeres II and III sepa- with an apical sclerite slender, recurved inwards, 
rate Lanyualtica from all other genera in the followed by a pair of longitudinal rows of short 
Tegyrius group. setae, then a pair of longitudinal rows of short teeth, 
This genus also resembles Psylliodes by the and with one elongate and laterally flattened basal 
similar body color, similar shape of the body sclerite.
and pronotum, and lacking supraantennal calli but Female. Length 2.88–3.27 mm, width 1.48–1.64 
is easily distinguished by its antenna with 10 mm. Antenna (Fig. 1B) similar to males, ratio of 
antennomeres. length of antennomeres I to XI 1.0:0.6:0.5:0.6:0.8:
Distribution. One species (described herein) 0.7:0.8:0.8:0.8:0.7:1.1; ratio of length to width 
known from Taiwan (Lanyu Island). of antennomeres I to XI 2.4:1.9:2.1:2.1:2.8:2.1: 
2.4:2.3:2.3:1.9:2.3. Sternite VIII (Fig. 1F) weakly 
Lanyualtica hsui Lee and Konstantinov, sclerotized, transparent, with dense, short setae 
new species along apical margin, several setae near apical mar-
zoobank.org/urn:lsid:zoobank.org:act: gin, spiculum extremely long. Spermathecal recep-
78BD3528-5D50-4352-ADBB-1C4550DFA77C tacle (Fig. 3G) less swollen; pump long and strongly 
(Figs. 1–3) curved, apically narrowed and apex narrowly round-
Types. Holotype ♂ (TARI): TAIWAN. Taitung ed; spermathecal duct with long basal part, ramus 
County: Lanyu Island (蘭嶼), Weather Station rounded, duct long. Vaginal palpi (Fig. 3H) slender 
(氣象站), 20.III.2023, leg. Y.-F. Hsu. Paratypes: 2♂, and conjoined for short distance at base, each gono-
7♀ (1♂, 7♀: TARI; 1♂, 2♀: USNM), same data as coxa apically narrowed, apex narrowly rounded, 
holotype; 3♀ (TARI), same but with “17.VI.2023”; with nine pairs of long setae along apex and lateral 
1♂, 5♀ (BMNH), same locality, 327 m, N 22º02.238 margin at apical one-fourth.
E 121º33.487, 26.VII.2008, hand collecting, leg. M. Etymology. The species is named after Dr. Yu-
V. L. Barclay and H. Mendel; 1♂ (USNM), 1♀ Feng Hsu (徐堉峰), a Taiwanese butterfly specialist 
(TARI), same island, East 81 Road, 14.III.2023, leg. who collected the type series of this species.
Y.-Y. Lu and Y.-F. Hsu. Host Plants. Chionanthus ramiflorus Roxb. 
Description. Color black (Figs. 1A–F), mouth- (Oleaceae). Adults of this new species were collect-
parts and prothoracic venter dark brown, meso- and ed together with another undescribed species of flea 
metathoracic venter and abdominal ventrites black- beetles (Argopistes Motschulsky sp.).
ish brown; antennae and legs brown with darker Distribution. Taiwan (Lanyu Island).
metafemur. Pronotum convex, with coarse, well- 
defined punctures; disc with reticulate microsculp- DISCUSSION
ture. Elytra 1.5 times longer than wide, disc with 
punctures larger, but less defined than those on In addition to the six endemic chrysomelid 
pronotum, arranged into 11 longitudinal rows; with species listed in the Introduction, 13 additional spe-
fine punctures in between. Proportionate lengths cies have been recorded on Lanyu Island. These 
of metafemur, metatibia, metatarsomere I, metatar- include one chrysomeline: Phola octodecimguttata 
somere II, metatarsomere III, and metatarsomere (Fabricius, 1775) (Chûjô 1958; Lee and Geiser 
IV equal to 1:0.63–0.74:0.39–0.42:0.11–0.14:1.10– 2023); three galerucines: Aulacophora analis 
0.11:0.14–0.25. (Weber, 1801), Aulacophora indica (Gmelin, 1790), 
Male. Length 2.73–2.83 mm, width 1.39–1.41 and Aulacophora lewisii Baly, 1866 (Lee and 
mm. Antenna filiform (Fig. 1A), reaching middle Beenen 2015); three flea beetles: Altica birmanensis 
of elytron when laid over pronotum; antennomeres (Jacoby, 1896) (= Haltica cyanea Weber, 1801, re-
I and II broad, III narrowest, III to VIII gradually corded by Chûjô 1936; Suenaga and Lee 2023), 
Downloaded From: https://bioone.org/journals/The-Coleopterists-Bulletin on 25 Mar 2024
Terms of Use: https://bioone.org/terms-of-use	Access provided by Smithsonian Libraries and Archives
 THE COLEOPTERISTS BULLETIN 78(1), 2024 75
Nonarthra chengi Lee, 2014 (= Nonarthra variabilis VIII. Subfamily Eumolpinae. The Philippine 
Baly, 1862, recorded by Chûjô 1935a; Lee 2014), Journal of Science 85: 1–180.
and Sinocrepis obscurofasciata (Jacoby, 1892) Chûjô, M. 1958. A taxonomic study on the Chrysomeli-
(= Sinocrepis micans Chen, 1933, recorded by dae (Insecta, Coleoptera) from Formosa, Part X. 
Chûjô 1935b); three eumolpines: Chrysopida murina Subfamily Chrysomelinae. Quarterly Journal of 
Baly, 1867 (Chûjô 1956), Colasposoma viridicoe- the Taiwan Museum 11: 1–85.Duckett, C. N., and J. D. Daza. 2004. A new species of 
ruleum Motschulsky, 1860 (= Colasposoma ober- flea beetle in the genus Alagoasa Bechyné (Cole-
thueri Jacoby, 1896, recorded by Chûjô 1956), optera: Chrysomelidae: Galerucinae) from high-
and Rhyparida sakisimensis Yuasa, 1930 (Kimoto land Costa Rica. The Coleopterists Bulletin 58(1): 
1974); one criocerine: Lema trivittata Say, 1824 84–96. doi.org/10.1649/608.
(Lee and Matsumura 2013); and two cassidines: Kimoto, S. 1974. Notes on the Chrysomelidae from Tai-
Aspidimorpha furcata (Thunberg, 1789) and wan VII. Entomological Review of Japan 26: 
Cassida circumdata Herbst, 1790 [= Metriona triv- 21–26.
ittata (Fabricius, 1801)] (both species recorded by Lee, C.-F. 2014. Review of the genus Nonarthra Baly 
Chûjô 1934). In this study, two species, neither of (Coleoptera: Chrysomelidae: Galerucinae: Altici-ni) from Taiwan and Japan, with descriptions of 
which have been previously recorded from the two new species. Japanese Journal of Systematic 
island, were collected from a single plant species, Entomology 20(2): 251–263.
C. ramiflorus. This suggests that the true species Lee, C.-F., and R. Beenen. 2015. Revision of the genus 
richness on Lanyu Island may be underestimated. Aulacophora from Taiwan (Coleoptera: Chrysome-
lidae: Galerucinae). Zootaxa 3949: 151–190. doi.
org/10.11646/zootaxa.3949.2.1.
ACKNOWLEDGMENTS Lee, C.-F., and M. F. Geiser. 2023. Taxonomic review 
on the genus Phola Weise, 1890 (Coleoptera: 
We thank Chi-Lung Lee for photos of specimens. Chrysomelidae: Chrysomelinae). Journal of Tai-
This study was supported by the National Science wan Agricultural Research 72(3): 197–218. doi.
and Technology Council NSTC 112-2313-B-055- org/10.6156/JTAR.202309_72(3).0002.
001-MY3. Mention of trade names or commercial Lee, C.-F., and Y. Matsumura. 2013. On newly and 
products in this publication is solely for the purpose recently recorded species of the genus Lema 
of providing specific information and does not Fabricius (Coleoptera, Chrysomelidae, Criocerinae) 
imply recommendation or endorsement by the from Taiwan. ZooKeys 262: 17–37. doi.org/ 
USDA; USDA is an equal opportunity provider and 10.3897/zookeys.262.4152.
employer. Prathapan, K. D., and A. S. Konstantinov. 2021. Revi-sion of Oriental flea beetle genera with subparal-
lel intercoxal ridges on the first abdominal 
REFERENCES CITED ventrite (Coleoptera, Chrysomelidae, Galerucinae, Alticini). Journal of Natural History 55: 1521–
Chûjô, M. 1934. Studies on the Chrysomelidae in the 1598. doi.org/10.1080/00222933.2021.1933229.
Japanese empire (V). Sylvia 5: 1–38. Reid, C., L. Halling, and M. Beatson. 2021. Revision of 
Chûjô, M. 1935a. Studies on the Chrysomelidae in the Australopapuan and West Pacific species of 
Japanese empire (VIII). Subfamily Halticinae (1). plain pumpkin-beetles, the Aulacophora indica 
Transactions of the Natural History Society of species-complex (Coleoptera: Chrysomelidae: 
Formosa 25: 354–369. Galerucinae). Zootaxa 4931: 1–73. doi.org/ 
Chûjô, M. 1935b. Studies on the Chrysomelidae in Jap- 10.11646/zootaxa.4932.1.1.
anese empire (VIII). Subfamily Halticinae (3). Suenaga, H., and C.-F. Lee. 2023. Taxonomic studies 
Transactions of the Natural History Society of on the genus Altica Geoffroy (Coleoptera, 
Formosa 25: 459–476. Chrysomelidae, Galerucinae, Alticini) in Taiwan: 
Chûjô, M. 1936. Studies on the Chrysomelidae in the Re-evaluation of A. himalayensis (Chen), A. cya-
Japanese empire (VIII). Subfamily Halticinae (4). nea (Weber), and A. caerulea (Olivier). Formosan 
Transactions of the Natural History Society of Entomologist 43: 49–65.
Formosa 26: 15–30.
Chûjô, M. 1956. A taxonomic study on the Chrysomeli- (Received 12 September 2023; accepted 19 January 
dae (Insecta: Coleoptera) from Formosa, Part 2024. Publication date 22 March 2024.)
Downloaded From: https://bioone.org/journals/The-Coleopterists-Bulletin on 25 Mar 2024
Terms of Use: https://bioone.org/terms-of-use	Access provided by Smithsonian Libraries and Archives