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SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY • NUMBER 518

Phylogenetic Relationships of Hedgehogs and Gymnures (Mammalia: Insectivora: Erinaceidae)

Darrel R. Frost, W. Chris Wozencraft, and Robert S. Hoffmann

SMITHSONIAN INSTITUTION PRESS

Washington, D.C.

1991

A B S T R A C T

Frost, Darrel R., W. Chris Wozencraft, and Robert S. Hoffmann. Phylogenetic Relationships of Hedgehogs and Gymnures (Mammalia: Insectivora: Erinaceidae). Smithsonian Contributions to Zoology, number 518, 69 pages, 12 figures, 19 plates, 8 tables, 1991.—Using 81 transformation series containing 172 characters of osteology, dentition, and external morphology, the phylogeny of Recent erinaceids is investigated using standard phylogenetic techniques. For this analysis, we used as the first taxonomie outgroup of Erinaceidae, Soricoidea + Tenrecoidea; and as the second taxonomie outgroup, †Leptictidae. Supraspccific monophyletic groups identified from this analysis are Erinaceidae; Hylomyinae; Podogymnura (aureospinula, truei); Echinosorex + Podogymnura; Hylomys (sinensis, suillus, hainanensis); Erinaceinae; Hemiechinus; subgenus Paraechinus ((aethiopicus, micropus), hypomelas) + H. auritus + H. collaris; Mesechinus dauuricus + (Erinaceus + Atelerix); Erinaceus (europaeus, concolor, amurensis); and Atelerix ((albiventris, sclateri), algirus, frontalis). Evidence does not support the monophyly of Hemiechinus (as recognized by Honacki et al., 1982) or of Hemiechinus excluding H. dauuricus. Mesechinus is elevated from subgeneric status (under Hemiechinus) to generic status to contain dauuricus and hughi. Paraechinus is considered a subgenus of Hemiechinus. Neohylomys and Neotetracus are considered synonyms of Hylomys. Diagnoses and taxonomie accounts of genera are presented along with comments on the taxonomy of and variation within species. Hemiechinus aethiopicus, H. deserti, and H. dorsalis are provisionally regarded as subspecies of H. aethiopicus.

OFFICIAL PUBLICATION DATE is handstamped in a limited number of initial copies and is recorded in the Institution's annual report, Smithsonian Year. SERIES COVER DESIGN: The coral Montastrea cavernosa (Linnaeus).

Library of Congress Cataloging-in-Publication Data

Frost, Darrel R.

Phylogenetic relationships of hedgehogs and gymnures (Mammalia: Insectivora: Erinaceidae) / Darrel R. Frost, W. Chris Wozencraft, and Robert S. Hoffmann.

p. cm.—(Smithsonian contributions to zoology; no. 518)

Includes bibliographic references.

1. Erinaceidae-Classification. 2. Erinaceidae-Morphology. 3. Phylogeny. I. Wozencraft, Wallace Christopher. II. Hoffmann, Robert S. III. Title. IV. Series.

Ql1.S54no. 518

[QL737.153]

591 s—dc20

[599.3′3] CIP

The paper used in this publication meets the minimum requirements of the American National Standard for Permanence of Paper for Printed Library Materials Z39.48–1984.

Contents

Introduction

Acknowledgments

Methods and Materials

Character Transformation Series

Cranial Characters

Dentition Characters

Post-cranial Characters

Non-skeletal Characters

Characters Not Utilized in Analysis

Results and Discussion

Recommended Taxonomy

Comparisons with Previous Hypotheses of Relationships

Key to the Skulls of the Erinaceidae

Taxonomic Accounts, Diagnoses, and Comments on Species Relationships

Family ERINACEIDAE

Subfamily HYLOMYINAE

Genus Echinosorex de Blainville, 1838

Genus Podogymnura Mearns, 1905

Genus Hylomys Müller, 1839

Subfamily ERINACEINAE Fischer, 1817

Genus Hemiechinus Fitzinger, 1866

Subgenus Paraechinus Trouessart, 1879

Genus Mesechinus Ognev, 1951

Genus Erinaceus Linnaeus, 1758

Genus Atelerix Pomel, 1848

Appendix 1: List of Specimens Examined

Appendix 2: Data Matrix

Appendix 3: Apomorphy List by Stems

Appendix 4: Changes on Stems by Transformation Series

Literature Cited

Plates 1–19

Requirements for Smithsonian Series Publication

Phylogenetic Relationships of Hedgehogs and Gymnures (Mammalia: Insectivora: Erinaceidae)

Barrel R. Frost, W. Chris Wozencraft, and Robert S. Hoffmann

____________

Darrel R. Frost, Museum of Natural History, The University of Kansas, Lawrence, Kansas 66045 (current address: Department of Herpetology and Ichthyology, American Museum of Natural History, Central Park West at 79th Street, New York, New York 10024); W. Chris Wozencraft and Robert S. Hoffmann, National Museum of Natural History/Office of the Assistant Secretary for Research, Smithsonian Institution, Washington, D.C. 20560.

Review Chairman: Stanley H. Weitzman, Smithsonian Institution. Reviewers: Gordon B. Corbet, British Museum (Natural History), John R. Wible, University of Louisville, and two anonymous reviewers.

Introduction

Erinaceomorphs are known from the lower Eocene (Novacek et al., 1985) with characteristics of the family already well established by the Early Oligocene (Butler, 1948). The Erinaceidae is a relatively plesiomorphic family of lipotyphlan insectivores that has its Recent distribution in the tropical and temperate zones of Eurasia and Africa. Current taxonomy divides the Recent members of the family into two nominally monophyletic subfamilies (Butler, 1948,1956,1988; Novacek, 1985, 1986; Corbet, 1988), Hylomyinae (= Galericinae = Echinosoricinae) (with 3–5 genera and 6 species) of the Southeast Asian tropics, and Erinaceinae (with 3–5 genera and 14 species), of Africa, Europe, and arid regions of Asia (Honacki et al., 1982; Nowak and Paradiso, 1983; Corbet, 1988). For purposes of discussion, we will use the nomenclature of Corbet (1988) (see Table 7), that is, 5 genera of hylomyines and 4 genera of erinaceines.

Because of their important position in larger questions of insectivore classification, erinaceids have been the object of considerable previous investigation. Leche (1902), Butler (1948, 1956, 1988), and McDowell (1958) studied the anatomy, and Novacek (1986) reviewed the relationships of the subfamilies within the framework of larger questions of erinaceomorph relationships. In an attempt to resolve the controversy surrounding generic distinctions in erinaceines, Robbins and Setzer (1985) evaluated the phenetic similarities of the erinaceine genera. More recently, Corbet (1988) reviewed the taxonomy, phytogeny, ecology, and zoogeography of Erinaceidae.

What has not been attempted is a rigorous approach to inferring the phylogenetic relationships of extant taxa of erinaceids from Recent material. In this paper we attempt such a phylogenetic analysis. More specifically, the questions asked are: (1) Are all of the nominal taxa within Erinaceidae monophyletic? (2) What is the most parsimonious hierarchical arrangement (= hypothesized historical relationships) of these discovered monophyletic taxa? (3) What taxonomy best reflects our knowledge of the relationships of these taxa?

Although the recent paper by Corbet (1988) might be considered an example of what we are trying to accomplish, we have reservations about his phylogenetic methods and taxonomy. Additionally, we could not verify some of the characters he used (see below). Because comparisons between our study and that of Corbet (1988) are unavoidable, we will highlight differences between our approaches, both philosophically and methodologically.

ACKNOWLEDGMENTS.—We are grateful to the curators and collection managers of the various museums who provided us with access to specimens used in this study; among them are Field Museum of Natural History (R. Izor, B. Patterson, L. Heaney); National Museum of Natural History (R. Fisher, A. Gardner, M. Carleton, L. Gordon, C. Handley, R. Thorington, D. Wilson); American Museum of Natural History (S. Anderson, K. Koopman, M. Lawrence, G. Musser); Harvard University Museum of Comparative Zoology (J. Kirsch, M. Rutzmoser); Academy of Natural Sciences of Philadelphia (C. Smart); Beijing, Institute of Zoology (Wang Sung); British Museum (Natural History) (I. Bishop, G. Corbet, J. Hill, J. Ingles, P. Jenkins); Delaware Museum of Natural History; University of Kansas, Museum of Natural History (R. Timm, T. Holmes); Moscow State University Zoological Museum (V.A. Dolgov, I.Ya. Pavlinov, O.L. Rossolimo). S. Sanford of Medsciart Co. did an excellent job of illustrating our specimens. V. Krantz photographed the specimens and prepared the plates. Appreciable improvements to the manuscript were made by several anonymous reviewers, to whom we are indebted. We also benefited from reviews and discussions with G. Corbet, J.R. Wible, A. Biknevicius, L. Heaney, R. Kay, and M. Novacek. Support for this project was provided by the University of Kansas and its Museum of Natural History and the Smithsonian Institution (National Museum of Natural History and the Office of the Assistant Secretary for Research).

Methods and Materials

We employed outgroup comparison, independent polarization of transformation series, and formation of the most parsimonious explanation of the data, a cladogram (Watrous and Wheeler, 1981; Wiley, 1979). For analysis, 81 transformation series containing 172 characters of external morphology, dentition, and osteology were collected from specimens of erinaceids of all nominal species, except Hemiechinus hughi (of which we have only seen the unique skull-less holotype), Paraechinus intermedius, and P. nudiventris (see Appendix 1, “Specimens Examined”). We used species for the operational taxonomie units (OTU) of our phylogenetic analyses to avoid problems resulting from unsuspected genus-level paraphyly.

Characters were initially drawn both from the literature (e.g., Butler, 1948, 1956, 1988; McDowell, 1958; MacPhee, 1981; Novacek, 1985, 1986; Corbet, 1988) and from our own observations. For purposes of outgroup comparison with Recent taxa, the arrangement of Novacek (1986) was accepted; that is, Soricoidea (Soricidae + Solenodontidae + Talpidae) + Tenrecoidea (Tenrecidae + Chrysochloridae) was considered the first taxonomic outgroup (sensu Watrous and Wheeler, 1981). Novacek (1985) and Butler (1988) noted a number of fossil erinaceomorphs that would normally be the nearest presumptive outgroups of erinaceids. Although fossil erinaceomorphs were not utilized directly as outgroups for our analysis, published information was used, when available (e.g., Novacek, 1985), for some polarity decisions. A fossil taxon, †Leptictidae († denotes fossil taxon), was considered the second taxonomic outgroup for our analysis. This choice was made because of the extensive summary of anatomy available (Novacek, 1986), and because second outgroup possibilities among Recent mammals demonstrated such an array of variation as to make comparisons unhelpful. Because of the taxonomic diversity in Soricoidea and Tenrecoidea, only the plesiomorphic conditions are noted for those taxa in the data matrix.

Much of the following discussion revolves around decisions made concerning the determination of polarity or the manner in which morphological variation is described that might influence the character placement on the tree. We attempted to describe the morphology involved in each transformation series in such a manner that it would be as unambiguous as possible. Where a particular morphological feature could be broken down into many recognizable components, we did so, and only considered reuniting those components when the correlation of their distribution was perfect. After a pattern of variability was described, each OTU was categorized to one state of that pattern, usually by the assignment of a letter code that implied no polarity decision. Finally, after all OTU’s had been coded, the polarity was assigned to each letter code by comparison to the outgroups. Generally, when both first and second outgroups agree in the polarity assigned to a transformation series, our rationale for adopting a particular polarity will not be discussed. Although polarity decisions are discussed with some transformation series, we have included data lines in our matrix (Appendix 2) for the plesiomorphic conditions found in Soricoidea, Tenrecoidea, and †Leptictidae to help in the understanding of our polarity decisions. Rooting of the cladogram was determined by using a hypothetical ancestor deduced from the distribution of characters in the outgroups.

Some character transformations could not be polarized adequately, due to ambiguous outgroup comparison. These features were used in the analysis as unpolarized (i.e., additivity retained) or unordered transformations (i.e., additivity not retained); that is, no polarity of transformation was assumed except those determined by correlation with the remaining characters. These transformations cannot affect the rooting of the calculated tree, but do serve to enhance stem definition within the rooted tree. Characters for all transformation series are presented in the data matrix for all outgroups (Appendix 2). In the following list, in the polarized transformation series, zero (0) denotes the plesiomorphic character and one (1) and two (2) denote progressively more derived characters; where the character was initially drawn from the literature, we have included that reference, although we do not necessarily code or state the character in the same manner. In unordered transformations the particular order of integers carries no information. In unpolarized transformations, additivity is retained although the ancestral condition was coded as “unknown,” allowing the polarity of the transformation to be determined by correlation with the otherwise most parsimonious solution of the data.

Analysis of data was completed using the PAUP (Phylogenetic Analysis Using Parsimony) program, version 2.4.1 (Swofford, 1984). The data matrix was analyzed using the multiple parsimony (MULPARS) and global swapping options. Global swapping allows the program to search for more parsimonious trees by global (as opposed to “nearest neighbor”) swapping of branches. MULPARS allows the swapping procedure to be performed on all topologically distinct trees of a given length rather than the first found of any particular length. Although the default option (FARRIS), used for optimization of character placement, prefers reversal over convergence, by using the CSPOSS and BLRANGE options (which note ambiguous placement of characters), as well as comparing results with DELTRAN outputs (an option that prefers convergence to reversal), and taking into account the distribution of “unknown” character assignments, evidence corroborating stems could be restricted to characters of unambiguous placement.

We have not employed differential character weighting. This practice has been argued against convincingly by Patterson (1982) and Novacek (1986). In short, character weighting assumes that in the absence of evidence from congruence of character distribution, one can “know” how “plastic” (in an a posteriori sense) a particular transformation is. Additionally, a posteriori weighting assumes that there are discoverable “natural” classes of transformations. Although we admit that pretests of homology are grounded in perceptions of similarity, we deny that injecting assumptions about the undetected levels of non-homology can increase the rigor of the analysis. Basically, weighting removes any objective grounds for discussion of character incongruence. Recently, Corbet (1988) has approached the systematics of erinaceids using subjective weighting of characters (see below).

In a number of features our observations disagree with previous authors. Although earlier studies were usually careful, the specimens available were (and in some cases still are) few and sampling errors were common. More unfortunate has been the tendency for some authors to refer to published literature and not to specimens. Many features that looked promising in our literature search did not survive specimen examination; we have tried to point out these discrepancies in our discussions. Because many of these features did not play central roles in previous discussions of phylogeny or identification, we have tended to ignore these for brevity’s sake. Before the reader assumes that we have missed a character noted by earlier authors we suggest that specimens be examined. However, following our list of transformation series is a section, “Characters Not Utilized in Analysis,” in which we discuss some of the features that have figured prominently in earlier studies.

We did not attempt to exclude apparent synapomorphies of Erinaceidae or interesting autapomorphies of terminal taxa from the analysis. Although such inclusion elevates the consistency index (CI), the only concern this could raise would rest on the belief that the consistency index is a measure of topological stability, or that it allows comparison between data matrices. Both of these beliefs are unfounded. We have used the consistency index as it was intended, only as a measure for comparing trees derived from the same data matrix (Kluge and Farris, 1969).

Character Transformation Series

CRANIAL CHARACTERS

We follow the nomenclature of Jayne (1898), McDowell (1958), MacPhee (1981), and Novacek (1986).

1. Nasal, posteriormost extension: (0) posteriormost portion of the nasals anterior to the level of the antorbital rim; (1) posteriormost portion of nasals extends medial or posterior to the level of the antorbital rim. This transformation is most likely related to rostral length (Transformation Series #2); however, the relationship is not necessarily a direct one as shown by the difference between Neotetracus sinensis (Plate 5) and Hylomys suillus (Plate 6).

2. Rostrum, length (Butler, 1948): (0) long (42% or more of skull length); (1) short (35% or less of skull length). Rostrum length is measured from the nasolacrimal foramen to the premaxi1la midline. Although outgroup comparison is ambiguous, in that many tenrecoids and soricoids have relatively short rostra, fossil erinaceomorphs have relatively long rostra (Novacek, 1986). We are therefore confident that, within Erinaceidae, we have characterized the transformation correctly. Butler (1956), Rich and Rich (1971), and Novacek (1986) also considered a long snout to be plesiomorphic within this clade. Corbet (1988) conversely considered the short snout of the erinaceines to be primitive and distinguished between the length of the rostrum in Podogymnura and Echinosorex on one hand and Hylomys, Neotetracus, and Neohylomys on the other. Our measurements of these taxa do not support Corbet’s separation into two rostrum length classes (Table 1).

3. Rostrum, width: (0) rostrum wide, anterior incisors not closely adjacent; (1) rostrum narrow, anterior incisors closely adjacent. In the plesiomorphic condition there is a distinctive diastema between the anteriormost incisors (e.g., compare Hemiechinus auritus, Plate 15, with Erinaceus europaeus, Plate 13).

4. Palatine foramina, size (Butler, 1948; Novacek, 1986): (0) small; (1) anterior foramina elongated posteriorly; (2) anterior foramina elongated to include middle palatine foramina. There is no reason to assume that the palatal perforations found in many erinaceids are homologous with those found in marsupials (Butler, 1972). Corbet (1988) did not distinguish among the anterior, middle, and posterior palatine foramina, and referred to “palatal perforations” as being absent, small, or large. His transformation series and distribution of characters would coincide with our series.

TABLE 1.—A comparison of selected skull measurement ratios, mean (range), of hylomyines and Erinaceus europaeus to the character state groups of Corbet (1988).

5. Anterior palatine foramina, location: (0) at the maxilla/palatine suture; (1) anterior to the maxilla/palatine suture. Corbet (1988) drew attention to the presence of long grooves in the palate for the major palatine artery in those taxa that have the anterior palatine foramina located anterior to the palatine/maxilla suture. The major palatine artery emerges ventrally from the anterior palatine foramen and continues anteriorly along a groove in the maxillary. These grooves are associated with the forward placement of the foramina (our Character 5.1); their distinctiveness and length are correlated with the age of the individual. There are faint, nondescript grooves in those taxa that have the foramina located at the suture, and these run forward in the palate, although not as distinctly as in the taxa with condition 5.1. We could not verify on our specimens Corbet’s contrasting of Hylomys and Neotetracus with short grooves and Podogymnura and Echinosorex with long grooves and suspect that this is individually variable with age.

6. Infraorbital canal, position of the anterior opening (Butler, 1948; Rich and Rich, 1971; Novacek, 1986): (0) dorsal or posterodorsal to the P4-M1 area; (1) dorsal to the P3–P4 region. Although most taxa coded as having the plesiomorphic condition have the anterior opening above P4-M1, Podogymnura truei has the opening more posterior, dorsal to the M1–M2 region (see Plate 3). For purposes of this analysis this species was coded as having condition “0.” In many soricoids and tenrecoids the position of the infraorbital foramen is set even farther posterior. We have refrained from considering P. truei as having a condition other than “0,” simply because of the ambiguity of outgroup comparison. This transformation reflects the relative shortening of the rostrum; the correlation is not perfect, however, so this transformation was retained as distinct from Transformation Series #2 (rostrum length).

7. Antorbital fossa (Butler, 1948, 1956; Novacek, 1986): (0) present; (1) absent. The antorbital fossa serves for the origin of the snout muscles and is located lateral and posterior to the infraorbital foramen on the anterior surface of the zygoma. Butler (1956) noted some structural differences between the antorbital fossa of †Leptictis and those found in tenrecoids, soricoids, and erinaceomorphs. Without evidence to the contrary, however, we regard these features as homologous. Although antorbital fossae are variably absent among some soricoids and tenrecoids, we regard presence to be the plesiomorphic condition. Subadults and most adults of Echinosorex do not exhibit the presence of an antorbital fossa, although some older individuals may develop an indistinct shallow fossa. Corbet (1988:123) referred to “horizontal crests behind infraorbital foramen” in his analysis of the hylomyines. If we interpret his transformation series correcüy, he referred to the deeply excavated antorbital fossa on the zygoma. In Hylomys, Neotetracus, Neohylomys, and Podogymnura the excavation of this fossa is reflected in the two ridges that surround it and meet posteriorly. In the erinaceines the fossa is absent and in those Echinosorex where a shallow fossa is present, it lacks a distinct lower rim.

8. Antorbital (= prelacrimal) flange (Butler, 1948, 1956): (0) not developed, so the lacrimal foramen is visible from the lateral view; (1) developed to a degree that the lacrimal foramen is hidden from lateral view. Butler (1948, 1956) discussed this feature from the viewpoint of the direction of the opening of the nasolacrimal foramen. Because the orientation of the nasolacrimal duct is merely a reflection of the development of the antorbital (= prelacrimal) flange we have cast the description of this feature in that light.

TABLE 2.—Abbreviations of morphological features used in figures and legends.

ACM

Acromion

MET

Metacromion

ALI

Ali sphenoid

MPF

Middle palatine foramen

ANT

Antorbital fossa

NAF

Nasolabilis fossa

APF

Anterior palatine foramen

NPF

Nasopharynx fossa

BAS

Basisphenoid, tympanic wing

NSV

Neural spine of vertebrae

CBL

Condylobasal length

PAG

Promontory artery groove

C/c

Upper/lower canine

PAR

Paroccipital process

COF

Condylar foramen

PET

Petrosal, tympanic wing

DEN

Dens

PGF

Poslglenoid foramen

EAM

External auditory meatus

PLF

Posterior lacerate foramen

ECA

Ectotympanic, anterior process

P/p

Upper/lower premolar

ECT

Ectotympanic

PPF

Posterior palatine foramen

EPI

Epipterygoid process

PRO

Promontorium

EPR

Epitympanic recessus

SAG

Stapedial artery groove

ETF

Ethmoid foramen

SCA

Sinus canal, anterior opening

EUS

Eustachian tube

SFA

Stapedial foramen, anterior opening

FIB

Fibula (fused)

SMF

Stylomastoid foramen

FOE

Foramen ovale

SQU

Squamosal

GLN

Glenoid fossa

STM

Stapedius muscle fossa

INF

Incisive foramina

SUP

Suprameatal fossa

KEL

Postventral keel of axis

TIB

Tibial crest or flange

M/m

Upper/lower molar

TRA

Transverse process

MAL

Manubrium of malleus

TYM

Tympanohyal

MAS

Mastoid process of the petrosal

FIGURE 1.—The basicranium of Paraechinus hypomelas (ventral view). Left auditory bullae cut away. (Scale = 5 mm; key to abbreviations in Table 2.)

9. Maxilla, posteroventral process on zygoma: (0) absent; (1) present, small and indistinct; (2) present, large and distinct.

10. Jugal, size (Butler, 1956; Novacek, 1985, 1986): (0) large, reaches lacrimal; (1) smaller, does not reach lacrimal, reaches posteroventral process of zygoma; (2) vestigial, confined to lateral rim of zygomatic process of maxilla; (3) absent. Even though a large jugal is clearly plesiomorphic for mammals, “local” outgroup comparison is uninformative in this case; therefore, this transformation must be treated as unordered. According to Butler (1972), the general reduction of the jugal can be considered evidence of monophyly of the lipotyphlans.

11. Pterygoid/alisphenoid and epipterygoid processes (parapterygoid of Osborn and Helmy, 1980) (Novacek, 1986): (0) epipterygoid processes absent, alisphenoid not inflated; (1) epipterygoid processes present, alisphenoid not inflated; (2) epipterygoid processes present, alisphenoid inflated; (3) epipterygoid processes present, alisphenoid and epipterygoid processes inflated. In Paraechinus the hypotympanic cavity of the middle ear is extended anteriorly into the base of the pterygoid/alisphenoid complex (Figure 1); in P. aethiopicus alone, it is expanded even farther into the epipterygoid process (Plate 14).

12. Lacrimal/maxilla suture (Butler, 1956): (0) unfused and/or distinct in adults; (1) fused and indistinct in young adults.

13. Frontal, supraorbital processes (Butler, 1948): (0) absent or poorly defined and blunt; (1) sharp, readily identifiable processes present. Our characterization of this transformation series differs from Corbet (1988) only in considering Neohylomys to share the derived condition with Hylomys and Neotetracus, rather than treating it as plesiomorphic. Corbet reported that the process in Neohylomys was not as well developed as those in the other taxa. Our examination of the type series leads us to believe that the process in Neohylomys is developed as well as or better than the process in the other taxa.

14. Frontal, supraorbital foramen (Novacek, 1986): (0) present; (1) absent. The absence of these foramina in the tenrecoids and soricoids makes outgroup comparison ambiguous. This transformation was therefore treated as unpolarized in our analysis.

15. Frontal inflation (Butler, 1948): (0) no inflation; (1) variable degrees of inflation. Butler (1948:450) noted that in this feature, Paraechinus and Hemiechinus resemble hylomyines. We could not characterize the difference in the degree of inflation of the frontals between Paraechinus, Hemiechinus, and the remaining erinaceines because of the difficulty in measuring and the apparent amount of individual variation. However, see Transformation Series #29 (skull height). Related directly to this feature, Corbet (1988) distinguished among the hylomyines by interorbital width. Our measurements do not support his division of the hylomyines into three distinct groups (Table 1).

16. Parietal, anterior process (Butler, 1948): (0) absent or very weak; (1) extends anteriorly along the supraorbital rim to form the base of the supraorbital process. The parietal has an anterolateral process that forms part of the postorbital rim and participates in the formation of the supraorbital process.

17. Alisphenoid, anterior process (Butler, 1948): (0) absent; (1) present. This character, related to the location of the sphenopalatine foramen, is apparently involved with the shortening of the orbitotemporal region. The orbital wing of the alisphenoid has a narrow, fusiform anterior process that sometimes fuses with a similar posterior process from the maxilla. The degree of formation seems to be individually variable. With the formation of this “bridge,” a narrow, short shelf is formed between the sphenopalatine and sphenorbital foramina.

18. Alisphenoid, inferior stapedial foramen, anterior opening (Corbet, 1988): (0) stapedial artery emerges from the auditory bulla in a groove in the alisphenoid; (1) stapedial artery emerges from the auditory bulla in a foramen in the alisphenoid. The inferior ramus of the stapedial artery emerges from the hypotympanic sinus on the anterior edge of the auditory bullae, slightly ventral and lateral to the foramen ovale. In hylomyines, this occurs in a groove in the alisphenoid (e.g., Podogymnura, Figure 2). In the erinaceines, there is a completely or incompletely formed foramen (e.g., Hemiechinus, Figure 3; Paraechinus, Figure 1). Corbet (1988) distinguished E. amurensis and E. concolor from all other erinaceines by the incomplete development of the inferior stapedial foramen. However, specimens we have examined of these and other erinaceine taxa reveal both conditions occurring, although the incomplete condition may be more prevalent in E. amurensis than in any other erinaceine taxon, as pointed out by Corbet (1988).

19. Frontal, ophthalmic foramen (Butler, 1948): (0) joined with, or closely adjacent to, the ethmoid foramen; (1) the foramina widely separated from the ethmoid foramen (e.g., see Figure 12).

20. Orbitosphenoid, suboptic foramen (Butler, 1948; Novacek, 1986): (0) absent; (1) present, in medial wall of sphenorbital fissure; (2) present, anterior to sphenorbital fissure. The “0” condition is unknown in erinaceids, which exhibit conditions “1” and “2.” Novacek (1986) has hypothesized implicitly 0→l→2, but we refrain from accepting this without sufficient outgroup comparison, and have treated this transformation as unordered in our analysis.

21. Palatine, sphenopalatine foramen (= orbitonasal foramen of Butler, 1948; sphenoid foramen of Novacek, 1986): (0) anterodorsal or slightly posterodorsal to the palatine transverse torus; (1) decidedly posterodorsal to palatine transverse torus. This transformation is difficult to visualize because it depends upon what horizontal plane is used for the skull. Therefore, we do not distinguish between those that are slightly anterior or posterior to the bisecting torus line.

22. Palatal shelf and spine (Corbet, 1988): (0) posterior palatal shelf with well-developed spine; (1) posterior spine absent or vestigial. In most taxa the palate extends posteriorly over the pterygoids (= mesopterygoid fossa) beyond the palatine transverse torus. Thomas (1918), Butler (1948), Robbins and Setzer (1985), and Corbet (1988) considered Atelerix to have a mesopterygoid fossa distinctly longer (“broader” in their terminology) than other erinaceines. We find variation of this feature in erinaceines to be so extensive as to be difficult to characterize, and polarized only ambiguously in comparison with outgroups. At least one specimen of Echinosorex (USNM 145584) has a decidedly long mesopterygoid fossa. In general, however, it appears that erinaceines have longer mesopterygoid fossae than hylomyines. A detailed analysis of this feature would likely document that there is a gradual statistical increase in the length of the mesopterygoid fossa as one progresses through the cladogram from hylomyines through erinaceines, finding its greatest development in specimens of Atelerix. Moreover, in specimens we have examined, Atelerix is easily distinguishable from other erinaceines on the absence of a well-developed posterior palatal spine (= posterior nasal spine of Jayne, 1898).

23. Palatine, lateral fossa, anterodorsal to the palatine transverse torus: (0) absent; (1) present. Development of a lateral fossa behind the palatal torus may be a reflection of the large size reached by Echinosorex. Its presence and size are perhaps related to the development of the medial pterygoid muscle.

FIGURE 2.—The basicranium of Podogymnura aureospinula (ventral view). Left auditory bullae cut away. The reproduction of the eclotympanic ring is taken from Podogymnura lruei. (Scale = 5 mm; key to abbreviations in Table 2.)

24. Squamosal, zygomatic process (Butler, 1948, 1956): (0) not elevated posteriorly; (1) elevated posteriorly. In the derived condition, the zygomatic process of the squamosal has a distinctive dorsal curve in the lateral view. Even though tenrecoids and soricoids (except for talpids) lack complete zygomatic arches, short zygomatic processes of the squamosal are present in those taxa and this condition therefore can be evaluated.

25. Squamosal, postglenoid foramen: (0) separated from glenoid fossa by entoglenoid process; (1) not separated.

The following three characters concern the shape, composition, and size of the deep pocket dorsal (intra-tympanic) and lateral to the tympanum (see Figures 1, 3). In the erinaceines, the tympanum stretches across two different planes: the eclotympanic is at an acute angle to the horizontal base of the skull and forms the medial attachment; the squamosal/mastoid pocket rim is on the horizontal plane of me skull and forms the lateral attachment for the tympanum. We are referring to this squamosal/mastoid excavation as the “suprameatal” fossa for purposes of discussion; we are uncertain whether it is homologous to the extratympanic “suprameatal fossa” in †Leptictis as described by Novacek (1986) or the fossa referred to by the same name present in some carnivores (both intra-and extratympanic) (Schmidt-Kittler, 1981). The homology of the various fossae and pockets that are associated with the external auditory meatus, whether they be ventrolateral or dorsolateral to the tympanum, is unclear (Wyss, 1987; Wozencraft, 1989). Although these three transformation series are most certainly related, the correlation is not perfect; therefore, they are listed separately.

26. Squamosal, suprameatal fossa, composition: (0) not well defined (“absent”) or with mastoid and squamosal contributions nearly equal; (1) squamosal and mastoid contributions subequal to predominately mastoid. We have chosen to include those taxa with poorly developed or no fossa (“absent”) together as the presence or absence of this feature is covered in Transformation Series #27.

FIGURE 3.—The basicranium of Hemiechinus dauuricus (ventral view). Left auditory bullae cut away. (Scale = 5 mm; key to abbreviations in Table 2.)

27. Squamosal, suprameatal fossa, depth: (0) absent; (1) shallow; (2) moderately developed; (3) deep; (4) very deep, extends dorsal to glenoid fossa. Corbet (1988) noted the distinction between H. dauuricus and other Hemiechinus, as well as the very inflated nature of Paraechinus. Osborn and Helmy (1980) noted the derived nature of P. aethiopicus and P. micropus. This is a series relating to the increase in volume of the suprameatal fossa. The description of characters “1” through “3” may sound as though they are somewhat subjective; however, these characters are quite distinctive and not easily confused. Unfortunately, without volumetric measurements, one will have to rely on reference taxa for the relative nature of each character.

28. Squamosal, suprameatal fossa, shape: (0) “normal,” anterior and posterior borders widely separated; (1) “compressed,” anterior and posterior borders narrowly separated (e.g., compare Figures 1 and 3).

29. Parietal, relative height of skull (Butler, 1948): (0) parietals relatively higher than frontals; (1) frontals relatively higher than parietals. Elevation of the frontals in Erinaceus and Atelerix makes them appear to have more highly inflated frontal sinuses than in Hemiechinus and Paraechinus. See Transformation Series #15 (frontal inflation).

30. Basisphenoid, nasopharyngeal fossa (= nasopharyngeal pocket of Butler, 1948): (0) absent; (1) present. The occurrence of condition “1” in Tenrec does not argue that this feature is a synapomorphy of Tenrecidae + Erinaceinae. Tenrec is far removed from the base of tenrec phylogeny (Eisenberg, 1981). In the derived condition, the basisphenoid contribution to the bullae is formed in such a manner that both sides touch medially and are closely appressed to the alisphenoid/pterygoid region. This enclosure, anteromedial to the bullae, is further excavated into a spherical fossa or hollow. In Paraechinus and Hemiechinus the hollow is slightly more anterior than in Erinaceus and Atelerix.

31. Basisphenoid, inflation (Corbet, 1988): (0) absent; (1) present. The basisphenoid forms part of the ventral floor and anteromedial wall of the hypotympanic chamber; in the derived condition it is relatively enlarged (Figure 4A). This feature is related to Transformation Series #36, although the correlation is not perfect. It refers to the relative contribution and size of the ventral expansion of the hypotympanic chamber, which is composed of three elements, the relatively slender uninflated ectotympanic and the inflated basisphenoid and petrosal.

FIGURE 4.—Comparison of the relative inflation of the auditory bullae: A, Erinaceus amurensis; B, Hemiechinus auritus; and C, Mesechinus dauuricus, from the occipital aspect. (Scale = 10 mm.)

32. Ectotympanic (= tympanic of Corbet, 1988) (Butler, 1948): (0) slender, ring-shaped, loosely attached, with a small anterior process; (1) much broader, sometimes engulfing anterior process, firmly attached. Material of Podogymnura aureospinula from which to evaluate this feature was unavailable and is treated as “unknown” in the data matrix. Butler (1948) stated that Echinosorex lacks an anterior process on the ectotympanic, otherwise seen in Hylomys and Neotetracus in material he had at hand. In our material, this process is present in Echinosorex as in the other hylomyincs; in Echinosorex and Podogymnura truei the ectotympanic is partially hidden within the bulla, making observations difficult. Corbet (1988) separated the large hylomyines (Podogymnura, Echinosorex) from the small ones (Neotetracus, Hylomys, and Neohylomys) on the basis of the small genera having a relatively larger ectotympanic. We could not verify this distinction and believe the major difference is between the erinaceines and the hylomyines. Corbet (1988) also pointed out that Echinosorex and Podogymnura have weakly attached ectotympanics, easily supported in Echinosorex by the difficulty in finding specimens in collections with the bone still attached. However, our examination of Podogymnura truei showed an equal proportion of specimens having the ectotympanic present as in other hylomyines.

33. Petrosal: (0) Promontorium predominantly confined to bullar roof, squamosal does not participate in bullar roof; (1) Promontorium forms posteromedial wall, squamosal is major component of bullar roof (compare the left side of Figures 1 and 2). Depending on how one views the relative topology of this region, one could argue that the distinctiveness of these characters is either attributable to a posterior growth of the squamosal, or a shifting of the petrosal to the posteromedial wall of the bullae.

34. Petrosal, inflation of mastoid process in region between the exoccipital and squamosal (Heaney and Morgan, 1982): (0) absent; (1) present. The occipital face of the mastoid appears slightly inflated in the derived condition (see also Transformation Series #26).

35. Petrosal, mastoid portion of suprameatal fossa (Corbet, 1988): (0) not inflated; (1) inflated. The suprameatal fossa is composed of two bones, the squamosal and the mastoid portion of the petrosal. The relative contribution of these two elements varies (see Transformation Series #26); however, in Paraechinus, the fossa is greatly excavated (Figure 1). Corbet (1988) believed that the condition in P. aethiopicus is much larger than that found in other erinacines and that some Hemiechinus auritus resemble Paraechinus, a conclusion not supported by our specimens. However, there may be a subtle difference in the size of the fossa between H. auritus and other erinaceines.

36. Petrosal, expansion of ventral process (Corbet, 1988): (0) not inflated; (1) inflated. Corbet (1988) noted thai Hemiechinus auritus shares with Paraechinus the relative inflation of the petrosal component of the floor of the hypotympanic chamber. Although H. auritus does show considerable variation (perhaps greatest in H. a. megalotis), in the specimens that we examined, it does not approach the condition found in Paraechinus (see also Transformation Series #31).

37. Basioccipital/petrosal suture: (0) narrow slit, exposing the posterior portion of the inferior petrosal sinus; (1) basioccipital/petrosal suture closed; well-defined posterior lacerate foramen.

38. Petrosal, stapedial foramen (for the ramus superior stapedial nerve) (Butler, 1948): (0) posterior to squamosal/alisphenoid suture and posterior to postglenoid foramen; (1) located on squamosal/alisphenoid suture and close to the postglenoid foramen.

39. Epitympanic recess, lateral wall: (0) formed partially by squamosal; (1) formed entirely by mastoid.

40. Paroccipital process (Thomas, 1918): (0) small; (1) robust.

41. Exoccipital: (0) not expanded; (1) expanded. The exoccipital bulges outward, giving an “inflated” appearance in the cleaned skull. This transformation may be implicated in size-related shape changes. We do not have ontogenetic series with which to verify this hypothesis, so we have retained the naive transformational hypothesis. A number of soricoids have expanded exoccipitals, but we do not regard this as the plesiomorphic condition in those taxa.

42. Occipital condyle (Novacek, 1986): (0) condyle emarginated causing the condyle to appear slightly lobed; (1) no condylar emargination.

43. Basioccipital, condylar foramen location (Novacek, 1986): (0) anterior to ventral lip of condyle; (1) on anteroventral lip of condyle.

44. Mandible, coronoid process (Corbet, 1988): (0) narrow, pointed; (1) broad, rounded. Corbet (1988) grouped Podogymnura aureospinula with Hylomys as having an intermediate condition. We found no difference in the shape of the coronoid process between Hylomys and Neohylomys on one hand and the species of Podogymnura on the other.

DENTITION CHARACTERS

We follow the nomenclature of Woodward (1896), Leche (1902), and Butler (1948, 1956).

45. i1 (Leche, 1902; Butler, 1948): (0) present; (1) absent.

46. i1, relative size (Butler, 1948): (0) absent or small; (1) enlarged.

47. i2, relative size (Butler, 1948; Novacek, 1986): (0) nearly equal to other incisors; (1) greatly enlarged.

48. I2, size (Novacek, 1986): (0) greater than I3; (1) less than or equal to I3.

49. I3, number of roots (Butler, 1948; Robbins and Setzer, 1985; Corbet, 1988): (0) one root; (1) two roots, separate; (2) two roots, fused. Concomitant with the change in the number of roots, the I3 becomes larger than the I2. Because it is impossible to tell the polarity between characters “1” and “2,” we have treated this transformation as unordered. However, because outgroup comparison allows the inference that the ancestor of Erinaceidae had condition “0” we have thus coded the hypothetical ancestor. See Table 3 for summary of variation in erinaceine taxa.

50. C1, size (Butler, 1948; Corbet, 1988): (0) significantly larger than adjacent post-canine teeth; (1) slightly larger than adjacent post-canine teeth; (2) approximately equal in size to adjacent post-canine teeth. Corbet (1988) used the same transformation series but with polarity reversed.

TABLE 3.—Variation in number of roots of I3 and C1 in Erinaceus and Hemiechinus (totals <N are due to missing teeth).

51. C1, number of roots (Butler, 1948; Robbins and Setzer, 1985; Novacek, 1986; Corbet, 1988): (0) two roots; (1) one root or two roots fused. Novacek (1986) noted that two canine roots are primitive for eutherians and this is the condition in early erinaceomorphs. For this reason, we regard the 1-root condition in tenrecoids + soricoids and Erinaceus to be non-homologous.

We found this character to be geographically and individually variable (Table 3). However, all species of Erinaceus have members that exhibit upper canines with fused (or single) roots. Our hypothesis is that the common ancestor of Erinaceus was polymorphic for fusion of canine roots and that this polymorphism is retained in its descendants.

52. c1, relative size (Butler, 1948; Corbet, 1988): (0) approximately equal to, or smaller, than pl; (1) significantly larger than pl. The polarity of the transformation specified by outgroup comparison is surprising. Although enlarged canines is the widespread condition in mammals, local outgroup comparison supports our view that relatively small canines are plesiomorphic in erinaceids. The enlarged lower canines in Tenrec we regard as homoplastic. See following transformation series.

53. C1, relative size (Corbet, 1988): (0) equal to, or larger than, I3; (1) subequal to slightly smaller than I3. Neotetracus has apparently reduced the size of its canines from the primitive condition.

54. P1: (0) present; (1) absent. Because outgroup comparison is insufficient for polarization, this transformation series was treated as unpolarized in the analysis.

55. p1: (0) present; (1) absent.

56. P2, roots (Corbet, 1988): (0) two roots; (1) one root, or two roots well fused. Corbet (1988) reversed the polarity and considered our derived condition to be two characters; however, in the specimens we have examined, these features (one root and two roots fused) appear to be variable.

57. p3, roots (Butler, 1948; Novacek, 1985, 1986; Corbet, 1988): (0) two roots present, larger in size than p2; (1) one root present, nearly equal in size to p2; (2) absent. Corbet (1988) considered the same transformation series with reversed polarity.

58. P3, lingual lobe (Corbet, 1978, as P2): (0) present, well developed; (1) vestigial or absent. Outgroup comparison is insufficient for polarization, so this transformation was treated as unpolarized even though we think that this feature is related to Transformation Series #60. Butler (1948:461) reported that the P3 lingual lobe was even further enlarged in Atelerix frontalis and A. algirus (which he distinguished from A. albiventris and A. sclateri), but we have been unable to verify this observation in our material.

59. P3, size (Butler, 1948; Novacek, 1985; Robbins and Setzer, 1985): (0) normal; (1) reduced. Corbet (1978) referred to this tooth as P2. Butler (1948:461) noted variation in the number of roots across taxa; we have been unable to verify his observations.

60. P3, roots (Corbet, 1988): (0) three roots; (1) fewer than three roots. Corbet (1988) considered finer divisions of this transformation series, with the polarity reversed. Outgroup comparison supports our order; we found his characters (one root, two roots hardly separated, and two roots divergent) to vary interspecifically. This is related to the number of cusps (see Transformation Series #58) although the correlation is not perfect (e.g., in Podogymnura).

61. P4, hypocone (Butler, 1948; Novacek, 1985, 1986): (0) triangular, hypocone (if present) vestigial; (1) quadrate, hypocone present. Presumably, this feature is correlated with the size of P3. Novacek (1986) noted that the leptictid P5 (= traditional P4) has a “distinctly developed” hypocone; however, we cannot discern this in his illustrations, or in our material, and Novacek (1985:21, his table 7) indicated that a large, distinct hypocone is present or absent in other erinaceomorphs. Because of this outgroup ambiguity the transformation is treated as unpolarized.

62. P4, lingual roots (Butler, 1948): (0) one lingual root; (1) two unfused roots; (2) two lingual roots, fused. Butler (1948:461) reported the fused roots of erinaceines as a single root. Because outgroup comparison allows the inference that the ancestor of Erinaceidae had condition “0” we have coded the hypothetical ancestor thus, although the overall transformation is unordered.

63. Lower molars, trigonid (Novacek, 1985): (0) high trigonid (significantly higher than talonid), talonid short/vestigial; (1) low trigonid (nearly equal in height to talonid), talonid expanded, large.

64. M1, lingual roots (Butler, 1948): (0) separate; (1) fused for most of their length. Butler (1948:460) referred to the fused condition as a “single lingual root which is flattened bucco-lingually” he also noted that in Hylomys and Neotetracus the M1 hypocone had a posterior crest that follows the posterior edge of the crown. In our material we have been unable to see this feature.

65. M3, roots (Butler, 1948): (0) three roots; (1) two roots.

66. M3, hypocone (Butler, 1948; Novacek, 1985, 1986): (0) absent or weak; (1) present, well developed on buccal side. Although Butler (1948:460) used presence of a hypocone as a feature uniting his Echinosoricinae (= Hylomyinae), he stated the hypocone to be primitively present and lost in erinaceines. Out-group comparison does not support this view and presence was coded as derived. Butler (1948:460) also noted that the degree of the approximation of the M3 metacone and hypocone is less in Hylomys than in Echinosorex. Although we can see what he was referring to, across all taxa the feature becomes impossible to characterize.

67. M3, metacone (Butler, 1948): (0) present, well developed; (1) present, small; (2) absent.

68. m3, talonid: (0) present; (1) absent

POST-CRANIAL CHARACTERS

We follow the nomenclature of Jayne (1898) and McDowell (1958).

69. Axis, posteroventral keel: (0) absent; (1) present. The axis normally has a convex ventral surface. In the derived condition, there is a posteroventrally projecting flange midway between the transverse processes (Figure 5).

70. Axis, neural spine: (0) low; (1) high. Although this transformation is correlated with the previous transformation, we could not evaluate this condition in †Leptictis and have therefore regarded this transformation series as unpolarized. The normal condition is for the neural spine to be horizontally flattened along the dorsal edge with little or no slope to the anterodorsal edge. In the “1” condition (Figure 5) there is a long distinctive projecting spine.

71. Scapula, metacromion process (McDowell, 1958): (0) deltoid, amorphous projection; (1) long, fusiform projection. We have not treated the transformation as polarized in our analysis because we were unable to evaluate the condition in †Leptictis. The “0” condition is a rather amorphous projection with a deltoid general shape; the projection is approximately equal to, or only slightly larger than, the acromion process. The metacromion projects at an acute angle to the scapular spine. In the “1” condition (Figure 6) the process is fusiform and much longer than the acromion and at approximately 90 degrees to the scapular spine.

72. Sacral vertebrae, neural spines: (0) not fused into a continuous longitudinal plate; (1) fused into a continuous longitudinal plate. Because we could not evaluate the condition in †Leptictis we did not treat this transformation series as polarized in our analysis. In the “1” condition the neural spines of the sacral vertebrae fuse together to form one long continuous spine (Figure 7).

FIGURE 5.—Comparison of the axis vertebrae, lateral aspect, in Hylomyinae and Erinaceus. A, Erinaceus europaeus; B, Echinosorex gymnura; C, Neotetracus sinensis; D, Hylomys suillus; and E, Neohylomys hainanensis. (Scale = 5 mm; key to abbreviations in Table 2.)

FIGURE 6.—Comparison of the scapula, lateral aspect, in Hylomyinae and Erinaceus. A, Erinaceus europaeus; B, Echinosorex gymnura; C, Neotetracus sinensis; D, Hylomys suillus; and E, Neohylomys hainanensis. (Scale = 5 mm; key to abbreviations in Table 2.)

73. Ischium, posteroventral process (McDowell, 1958): (0) not greatly elongated; (1) greatly elongated. In the primitive condition, there is little, if any, posteroventral process to the ischium, the posteroventral border being rounded. In the derived condition, the posteroventral border (ischiatic tuberosity) is produced into long fusiform processes that projects a considerable distance from the obturator foramen (Figure 7).

74. Tibia, lateral flange: (0) absent or only weakly present; (1) lateral flange on antero-superior margin strongly developed. The tibial crest is produced into a distinctive lateral flange in the derived condition (Figure 8).

NON-SKELETAL CHARACTERS

75. External pinnae, length relative to condyloincisive length (Thomas, 1918; Corbet, 1974, 1988; and Robbins and Setzer, 1985, all used this character relative to body length): (0) short (considerably shorter than 65% of condyloincisive length); (1) long, (considerably longer than 65% of condyloincisive length) (Table 4). Even though leptictids cannot be evaluated for this feature, the uniqueness of this feature within lipotyphlan insectivores argues for long ears being apomorphic.

76. Hallux (Thomas, 1918; Robbins and Setzer, 1985; Corbet, 1988): (0) normal; (1) reduced, claw not reaching base of second digit; (2) absent. Because †Leptictis cannot be evaluated for this feature, other mammals were used as a functional outgroup. We were unable to evaluate this character in our specimen of Atelerix sclateri (skull only). However, Corbet (1988) stated that the hallux is present, though small.

FIGURE 7.—Comparison of the pelvic region, lateral aspect, in Hylomyinae and Erinaceus. A, Erinaceus europaeus; B, Echinosorex gymnura; C, Neotetracus sinensis; D, Hylomys suillus; and E, Neohylomys hainanensis. (Scale = 5 mm.)

77. Pelage, condition (Corbet, 1974, 1988): (0) normal, not spiny; (1) smooth spines; (2) spines papillate, not grooved; (3) spines papillate and grooved. We have relied heavily on Corbet (1988) for the character distribution of this feature. Although a few tenrecids have spiny pelage, the uniqueness of this feature within lipotyphlan insectivores argues for its being apomorphic, and Corbet (1988:131) asserted that the erinaceine spine “is very different from that of the superficially similar spines in the family Tenrecidae.” The character “spiny” is restricted to stout heavy spines. Podogymnwa aureospinula has been evaluated as having spiny pelage (Heaney and Morgan, 1982); although P. aureospinula does have well-developed guard hairs, they are only marginally stouter than those in Echinosorex and P. truei and do not approach the dagger-like development in erinaceines (Poduschka and Poduschka, 1985; Corbet, 1988). For this reason, P. aureospinula is coded as “0.” Because outgroup comparison is uninformative, this transformation is treated as unordered. Although “spines” is clearly a synapomorphy of erinaceines we have not included it, as such, as a character, for no other reason than “spines present” would be redundant with whatever the basal erinaceine shift from Character 77.0 would be.

78. Pelage, completely white spines among dark spines (Corbet, 1988): (0) absent; (1) present. Corbet (1988) noted that in certain erinaceines, there are distinctive completely white spines irregularly scattered among completely dark or banded spines. We confirmed his observation of the derived condition in E. amurensis and A. frontalis. Only about one-half of the specimens we examined of E. europaeus had the derived condition he indicated; therefore we coded it as “unknown.” Although Corbet (1988) also listed A. algirus as derived, we could not find specimens with this condition.

FIGURE 8.—Comparison of the tibia, lateral aspect, in Hylomyinae and Erinaceus. A, Erinaceus europaeus; B, Echinosorex gymnura; C, Neotetracus sinensis; and D, Hylomys suillus. (Scale = 5 mm.)

TABLE 4.—Measurements of the length of the external pinnae of erinaceines (CIL = condyloincisive length).

79. Pelage, medial tract on crown (Corbet, 1988): (0) no spines on top of head; (1) spineless medial crown tract narrow; (2) spineless medial crown tract wide; (3) spineless medial crown tract absent. Thomas (1918), Corbet (1974, 1988), and Robbins and Setzer (1985) recorded crown parting in all spiny genera except Hemiechinus, although they noted that it was sometimes inconspicuous. According to Corbet (1988:132), a spineless medial crown tract is present in “new-born Hemiechinus auritus,” suggesting that the adult condition is derived, although it could be that the adult condition in the other genera is neotenic; outgroup comparison does not shed light on this transformation. Because outgroup comparison is uninformative for the polarity of this transformation it was included in our analysis as unordered. This feature could not be evaluated in our material of Atelerix sclateri, but Corbet (1974, 1988) reported that a spineless crown tract is present.

80. Pelage, condition on ventral side (Corbet, 1974, 1984): (0) soft and densely furred; (1) coarse and relatively sparsely haired.

81. Pelage, body underfur (Corbet, 1988): (0) absent; (1) present. It is difficult in some specimens to determine the difference between underfur and very fine guard hairs. However, the taxa with the derived condition clearly have two types of hair, whereas in others only one is present.

Characters Not Utilized in Analysis

A number of features have been used in analyses by other authors, but for a number of reasons we have not utilized them here. These features, as well as interesting anatomical novelties that may prove useful to future workers, are discussed.

1. Maxillae, posterior processes: Corbet (1988) united Erinaceus on the shortening of the posterior rostral process of the maxilla. In Erinaceus, the nasolabilis fossa is only partially composed of the maxilla, with the posteriormost point of the maxilla at or near the level of the lacrimal foramen. We concur with Corbet in recognizing this tendency in Erinaceus, which is not found in other erinaceines; however, we found the character too variable within taxa to code.

2. Occipito-parietal suture: The contribution of the parietal bone to the occipital crest, or lack thereof, was used by Corbet (1974) to distinguish among African hedgehogs. We found this character too variable to be useful although, a posteriori, a trend is visible.

3. Nasal-maxilla relationship (Corbet, 1974, 1988): We concur with Leche (1902) and Butler (1948) and find that the contact between the nasal and maxilla exhibited too much intraspecific variation to be useful (Table 5).

TABLE 5.—A comparison of the nasal/maxilla suture length (mm) in the Erinaceinae to the groups of Corbet (1988).

4. p4, metaconid (= lingual cusp of Corbet, 1988): Corbet (1988) used this character to differentiate among erinaceines (Erinaceus and Atelerix with large cusp; Paraechinus and Hemiechinus with small cusp; and Hemiechinus dauuricus and Atelerix algirus with no cusp). We also found no cusp in the series of H. dauuricus that we examined. However, we found the variation within species much greater than indicated by Corbet, with all three conditions occurring in most taxa.

5. “Tympanic angle” in posterolateral margin: Corbet (1988) used this character to distinguish Hemiechinus dauuricus and Paraechinus from other erinaceids. He coded the tympanic angle in the margin as either acute or obtuse. We have not been able to verify this character on our specimens.

6. Mammae (Corbet, 1988): The number of mammae is difficult to evaluate from dry skins and we are skeptical of published values, because much variation is evident (Table 6).

7. Tail length (Corbet, 1988): Cursory examination of erinaceids shows considerable variation in tail length, from a very long tail in Echinosorex to the very short tail of erinaceines. If these extremes were all there were, we would have no trouble employing tail length as one of our characters. Podogymnura, Hylomys, Neohylomys, and Neotetracus are roughly intermediate but difficult to characterize, given the condition of the skins available to us.

8. Pelage, white facial mask: Corbet (1988) distinguished among erinaceines on the presence of a white facial mask. He listed Atelerix, Paraechinus aethiopicus, and P. micropus as having a distinct mask, and Erinaceus europaeus and E. concolor as having a faint mask. Although we agree that there may be a trend in this direction, we found the presence of a white mask too variable within these species to warrant coding.

9. Pelage, dark sagittal stripe: Corbet (1988) used this character to distinguish between Hylomys/Neotetracus with a faint stripe and Neohylomys with a prominent stripe. Many of the specimens that we have examined of Hylomys and Neotetracus have no evidence of a stripe, and the type series of Neohylomys shows considerable variation in the distinctiveness of the stripe.

TABLE 6.—Reported values for numbers of mammae in the Erinaceidae.

10. Pelage, ventral pigmentation: Corbet (1988) coded (he variation in pigmentation on the ventral surface in erinaceines. We found the color to vary individually, perhaps being more closely associated with environmental variables.

11. Pre-anal glands: Corbet (1988) used this character to separate Echinosorex with pre-anal glands on the anterior margin of the anus from Hylomys and Neotetracus, where the pre-anal glands are separate from the anal opening. Pre-anal glands appear to be absent in the few erinaceines checked and the condition is unknown in Podogymnura or Neohylomys. Because of the paucity of information, we have excluded this character.

12. Penis spines: Corbet (1988) pointed out the differences in the arrangement and nature of spines on the penis. We were not able to examine material and therefore did not include this transformation in our analysis. Note, however, that Corbet’s (1988) Figured distribution of penis spines is more congruent with our tree than his.

13. Penis, glans expanded laterally: Corbet (1988) pointed out that the expansion of the glans penis separated some erinaceines (A. algirus, A. albiventris, and A. frontalis) from the others. We were unable to examine material and therefore did not include his character in our analysis, although it supports our tree.

Results and Discussion

The least-conflicting global phylogenetic hypothesis (CI = 0.758; 128 steps) is presented in Figure 9. The characters supporting this tree are listed in Appendix 3 (Apomorphy List by Stems) and Appendix 4 (Changes on Stems by Transformation Series). The results of this analysis are as follows:

FIGURE 9.—Recovered phylogenetic tree. Lower numbers are the stem numbers (see text), upper numbers are the number of unambiguous apomorphies. Taxonomy used is that of Corbet (1988).

1. The monophyly of Erinaceidae is highly corroborated by seven unambiguously placed characters (Figure 9, Stem 1; Appendix 3, Stem 1).

2. Hylomyinae (Stem 2) is monophyletic, although without postcranial evidence this would not be clear. Some evidence (i.e., 4.1, 6.1, 50.1, 57.1, and 59.1) suggests that Hylomys, Neotetracus, and Neohylomys are most closely related to erinaceines. Other features, however, both cranial and postcranial (i.e., 8.1, 33.1, 66.1, 69.1, 73.1, and 74.1), support the monophyly of Hylomyinae.

3. The Hylomys group (Neotetracus, Neohylomys, and Hylomys) (Stem 5) is well supported by seven unambiguously placed character shifts, of which three features of the orbit are unique and unreversed (i.e., 13.1, 16.1, and 19.1). The relationship between the three species currently placed in the three monotypic genera is only arguably resolved. Although in our preferred cladogram, Hylomys and Neohylomys are joined by 54.0 (reappearance of the P1), this requires the ancestral erinaceid to have lost the P1, which then reappeared in Echinosorex and Hylomys + Neohylomys. There has been a traditional bias among mammalogists not to support the loss and then later reappearance of a homologous tooth. A one step less parsimonious solution, that of independent loss of the P1 in Podogymnura, Neotetracus, and erinaceines, remains a viable possibility, and would support the generic distinction of Neotetracus from Hylomys.

4. Podogymnura (Stem 4) is arguably monophyletic, supported by two unambiguously placed characters, 22.1 and 48.1. Although two cranial characters (34.1 and 41.1) argue that P. truei is more closely related to the Hylomys group than to P. aureospinula, for these to be homologues would require the unique unreversed apomorphies of Stem 4 and Stem 3 to be homoplasies. Also there is some evidence that the relative size of these features may be allometric.

5. A sister-taxon relationship of Echinosorex and Podogymnura (Stem 3) is supported by four unambiguously placed apomorphies (5.1, 44.1, 52.1, and 81.1). Additionally, the tendency for the ectotympanic to be somewhat hidden within the bulla supports this conclusion.

6. Erinaceinae (Stem 7) is highly corroborated with 30 unambiguously placed synapomorphies. Within this group, two groups can be recognized: (1) a well-supported clade composed of Hemiechinus (excluding H. dauuricus) and Paraechinus (Stem 8); (2) a weakly supported clade composed of Hemiechinus dauuricus, Erinaceus, and Atelerix (Stem 11).

7. The monophyly of the Hemiechinus (excluding H. dauuricus)-Paraechinus clade (Stem 8) appears strong (although disputed by Corbet, 1988), supported by three cranial and one external pelage character (3.1, 27.2, 31.1, and 75.1).

8. Paraechinus (Stem 9) is also a well-supported monophyletic group, supported by unreversed basicranial characters (11.2, 27.3, 35.1, and 36.1), as well as one unordered feature (79.2).

9. Within Paraechinus it is clear that Paraechinus hypomelas is the sister taxon of P. micropus + P. aethiopicus (Stem 10). Paraechinus hypomelas does not exhibit any discovered apomorphies with respect to Stem 10 and could therefore be considered to approach the diagnosis of the ancestor of living Paraechinus. Likewise, P. micropus lacks any apomorphies with respect to P. aethiopicus.

10. Hemiechinus is not monophyletic, because H. dauuricus demonstrates a possible special relationship with the Erinaceus-Atelerix clade (Stem 12). It may be argued that the degree of difference between Paraechinus (Stem 9; see point 8 above) and Hemiechinus (H. auritus and H. collaris, excluding H. dauuricus) warrants generic distinction. However, we have no evidence to support the monophyly of the remaining Hemiechinus and it remains possible that, regardless of overall similarity, Hemiechinus (sensu stricto) is paraphyletic with respect to Paraechinus.

11. The monophyly of the clade composed of Hemiechinus dauuricus, Erinaceus, and Atelerix (Stem 11) is weakly supported by Character 80.1 (coarse ventral pelage). Single characters make poor flagpoles, however, and we are not overly confident of this conclusion, because at least one other Transformation Series, #77 (spine condition), if considered an ordered series, would argue for the monophyly of H. dauuricus + (other Hemiechinus + Paraechinus). Hemiechinus dauuricus is best considered to be in a polytomy with the Hemiechinus -Paraechinus clade (Stem 8) and the Erinaceus-Atelerix clade (Stem 12).

12. The Erinaceus-Atelerix clade (Stem 12) is supported by two characters of the skull and pelage (29.1 and 79.1) the second of which is part of an unordered series.

13. Erinaceus (Stem 16) has its monophyly supported by two unique unreversed features of the teeth (49.2 and 51.1) and a reversal (76.0). In trees of one extra step in Transformation Series #80, Character 76.0 succeeds, in some topologies, in placing Erinaceus as the sister taxon of the remaining erinaceines. No resolution between the species of Erinaceus was obtained.

14. The Atelerix clade (Stem 13) is supported by three unambiguously placed characters of the skull and teeth (22.1, 26.1, and 56.0).

15. Aethechinus (sensu Robbins and Setzer, 1985; i.e., Atelerix algirus and A. frontalis) does not have any apomorphies to support an hypothesis of monophyly. Conversely, however, Character 60.1 (P3 with two roots) supports paraphyly of Aethechinus (Stem 14). Additionally, even though we could not use the shape of the parieto-occipital suture (Corbet, 1974) because of individual variation, the “trend” suggests the same results: that Aethechinus is, in fact, paraphyletic, with A. algirus being closer to Atelerix than is Aethechinus frontalis.

16. Atelerix albiventris plus A. sclateri (Stem 15) is monophyletic, supported by one feature on P3 (58.1).

Recommended Taxonomy

Although any number of taxonomies are consistent with our preferred tree (Figure 10), we have designed a taxonomy that attempts to maintain usage without distorting phylogenetic information. Our adopted taxonomy is listed below. Salient features are as follows:

1. Following Van Valen (1967) and Nowak and Paradiso (1983), the monotypic genera Neohylomys and Neotetracus are synonymized with Hylomys. We could continue to recognize three monotypic genera (as did Corbet, 1988), but that arrangement would carry no phylogenetic information and would leave the highly corroborated stem subtending these taxa without a name.

2. Hemiechinus dauuricus (and tentatively, H. hughi) is placed in Mesechinus, which we regard as in a polytomy with the Hemiechinus-Paraechinus clade and the Erinaceus-Atelerix clade.

3. Paraechinus is placed as a subgenus within an expanded Hemiechinus. Although we have no evidence bearing on the monophyly of a subgenus Hemiechinus, it appears that Hemiechinus collaris may be a peripheral isolate of H. auritus and, therefore, would form a monophyletic taxon. Further data are needed.

4. Following earlier authors (Table 7), species formerly in Erinaceus, Aethechinus, and Atelerix are placed in two genera, Erinaceus and Atelerix (including former Aethechinus). Although a phylogenetically more symmetrical taxonomy would have Atelerix and Erinaceus as sister subgenera, the evidence linking these two taxa is only one unambiguously placed character. Because trees of one step longer allow Erinaceus to form the sister taxon of the remaining genera, we maintain the two genera. Regardless, our taxonomy is logically consistent with our recovered phylogeny.

Where the species listed differ from those discussed previously, see the relevant taxonomic accounts. Comparison of our taxonomy with those used by other authors is shown in Table 7. The taxonomy we propose follows.

Family ERINACEIDAE

Subfamily HYLOMYINAE

Hylomys

H. hainanensis

H. sinensis

H. suillus

Echinosorex gymnura

Podogymnura

P. aureospinula

P. truei

Subfamily ERINACEINAE

Hemiechinus

H. auritus

H. collaris

H. (Paraechinus)

H. (P.) aethiopicus

H. (P.) hypomelas

H. (P.) micropus

H. (P.) nudiventris

Mesechinus

M. dauuricus

M. hughi

Erinaceus

E. amurensis

E. concolor

E. europaeus

Atelerix

A. algirus

A. frontalis

A. albiventris

A. sclateri

FIGURE 10.—Taxonomic tree reflecting our proposed nomenclature.

TABLE 7.—A comparison of classifications of Recent Erinaceidae.

Honacki et al., 1982

Corbet, 1988

This paper

Echinosorex gymnurus

Echinosorex gymnurus

Echinosorex gymnura

Podogymnura aureospinula

Podogymnura aureospinula

Podogymnura aureospinula

Podogymnra truei

Podogymnura truei

Podogymnura truei

Hylomys suillus

Hylomys suillus

Hylomys suillus

Neohylomys hainanensis

Neohylomys hainanensis

Hylomys hainanensis

Neotetracus sinensis

Neotetracus sinensis

Hylomys sinensis

Erinaecus amurensis

Erinaecus amurensis

Erinaecus amurensis

Erinaecus concolor

Erinaecus concolor

Erinaecus concolor

Erinaecus europaeus

Erinaecus europaeus

Erinaecus europaeus

Erinaceus albiventris

Atelerix albiventris

Atelerix albiventris

Erinaecus frontalis

Ateleeix frontalis

Atelerix frontalis

Erinaceus algirus

Atelerix algirus

Atelerix algirus

Erinaceus sclateri

Atelerix sclateri

Atelerix sclateri

Hemiechinus auritus

Hemiechinus auritus

Hemiechinus auritus

Hemiechinus collaris

Hemiechinus collaris

Hemiechinus collaris

Hemiechinus dauuricus

Hemiechinus dauuricus

Mesechinus dauuricus

Hemiechinus hughi

Hemiechinus hughi

Mesechinus hughi

Hemiechinus sylvaticus

(Hemiechinus hughi)

(Mesechinus hughi)

Paraechinus aethiopicus

Paraechinus aethiopicus

Hemiechinus aethiopicus

Paraechinus hypomelas

Paraechinus hypomelas

Hemiechinus hypomelas

Paraechinus micropus

Paraechinus micropus

Hemiechinus micropus

Comparisons with Previous Hypotheses of Relationships

Other than Leche (1902), most authors (e.g., Butler, 1948; Novacek, 1985, 1986) have accepted the monophyly of Hylomyinae (as Galericinae or Echinosoricinae), albeit without much supporting data. Leche (1902) supported the view that Hylomys had a special relationship with the erinaceines; without postcranial data this view would be as parsimonious as one supporting monophyly of the hylomyines.

Heaney and Morgan (1982) and Corbet (1988) considered Podogymnura to be a monophyletic taxon, most closely related to Echinosorex. The former presented no evidence to support this conjecture, other than a subjective evaluation of general similarity, but Corbet (1988) presented a number of features that were used in our analysis. Butler (1948) also regarded Podogymnura to be most closely related to Echinosorex (together in a tribe, Echinosoricini), based on overall similarity.

Although the monophyly of Erinaceinae has been universally accepted, the number of erinaceine genera has been the source of some controversy. Dobson (1882) considered all erinaceines to be in Erinaceus. Corbet (1974, 1978) accepted three genera, Erinaceus (including Atelerix), Hemiechinus, and Paraechinus. Ellerman and Morrison-Scott (1951) and Nowak and Paradiso (1983) recognized the same three genera, but recognized two subgenera, Erinaceus and Atelerix, within the genus Erinaceus. Corbet (1988) later separated Atelerix from Erinaceus. With some disagreement on the assignment of some species, Thomas (1918), Cabrera (1925), Allen (1939), and Robbins and Setzer (1985) accepted five genera: Atelerix, Aethechinus, Erinaceus, Hemiechinus, and Paraechinus. With the exception of the one-genus arrangement of Dobson (1882), which does not improve on the information implicit in the group name Erinaceinae, none of these arrangements is consistent (Hull, 1964; Wiley, 1981) with our inferred phylogeny of the group. That is, they recognize at least one paraphyletic genus, Hemiechinus.

Most of these authors based their genera only on some subjective measure of similarity rather than relationship. Only Robbins and Setzer (1985) supplied a comprehensive rationale for their arrangement, but they had conflicting goals (e.g., recognize monophyletic taxa, group phenetically similar species, and identify “adaptive zones”). Their scheme of relationship is almost completely in disagreement with ours. In their dendrogram Aethechinus and Erinaceus are allied, apparently solely on the basis of size, as there were no synapomorphies suggested, nor have we found any support for this hypothesis. We also know of no support for the monophyly of their group containing Atelerix (in the more restricted sense), Hemiechinus (in the sense of including dauuricus), and Paraechinus. Our only point of agreement is the close relationship of Paraechinus and Hemiechinus (excluding dauuricus). We, however, do not regard their Hemiechinus as demonstrably monophyletic. Our differences from the evolutionary model presented by Robbins and Setzer (1985) appear to be due to the fact that phenetic techniques, because of their underlying assumptions, are inappropriately applied to historical questions (Sokal, 1986).

Most recently, Corbet (1988) has addressed the systematics of Erinaceidae using techniques superficially similar to ours and has come to a number of different conclusions. The reason for these differences stems from his use of subjective weighting. Although we do not categorically state that his approach renders his conclusions untestable, we do think that subjective weighting does not allow for objective comparisons with our results. Corbet (1988) regarded Hylomys suillus as the sister taxon of Neohylomys (our Hylomys hainanensis) and Neotetracus (our H. sinensis) on the basis of their shared loss of the P1. However, the presence of this tooth is variable in the type series (Shaw and Wong, 1959). One more parsimonious solution is that the P1 reappeared in Hylomys hainanensis + H. suillus. Although Corbet (1988) retained the monotypic genera Neohylomys and Neotetracus as separate from Hylomys, this difference has nothing to do with data analysis, but rather with a difference of opinion as to the importance of phenotypic similarity. We see no value in recognizing monotypic genera when to do so does not improve perception of cladistic relationships.

In the erinaceines, Corbet (1988) regarded Hemiechinus (including dauuricus) as the sister taxon of the remaining erinaceines on the basis of not having a spineless tract on the scalp. Regardless of the attractiveness of this notion, it requires at least five convergences between Paraechinus and Hemiechinus, excluding dauuricus (i.e., 3.1 (incisors closely approximating medially), 27.2 (moderately developed suprameatal fossa), 31.1 (basisphenoid inflation), and 75.1 (enlarged ears)), and also requires the coarse ventral fur (Character 80.1) to be convergent in former Hemiechinus dauuricus and the Erinaceus-Atelerix group. We believe it inappropriate to base the tree on a single transformation (presence/absence of a spineless crown tract), particularly when this transformation cannot be polarized.

Another relationship suggested by Corbet (1988) was the alliance of Paraechinus with Atelerix, radier than with Hemiechinus (excluding dauuricus), a result inconsistent with our tree. When compared with our data, each of his proposals necessitates considerably more homoplasy. As a general observation, when our data are cast on Corbet’s tree, the obtained consistency index is 0.703 (compared with 0.758 for our tree). Although these numbers appear similar, bear in mind that 25% of all character shifts in our analysis appear on the Erinaceinae stem. What these numbers do reflect is a lack of parsimony in Corbet’s tree. We concur, however, with his species limits.

Key to the Skulls of the Erinaceidae

Taxonomic Accounts, Diagnoses, and Comments on Species Relationships

All characters supporting the tree in Figure 9 are supplied in Appendices 3 (list of character changes by stem) and 4 (list of changes by transformation series). For this reason, with the exception of Erinaceidae, the diagnoses supplied with the accounts are not lists of apomorphics. Instead they are comparable lists of attributes that will serve to allow easy comparison. Numbers in brackets indicate character transformation series. Synonymies include names based on fossils only when those names affect the stability of current nomenclature (e.g., Galericini). Type localities are listed with each species. When possible, we have recorded the corresponding longitude and latitude of the type locality as listed in the U.S. Board of Geographic Names Gazetteers. This information follows the type locality description.

Family ERINACEIDAE Fischer, 1817

ERINACEIDAE Fischer, 1817:372. [Type genus: Erinaceus Linnaeus, 1758.]

DIAGNOSIS.—Erinaceidae is apomorphic with respect to soricoids, tenrecoids, and †Leptictidae in having (1) an epiptcrygoid process of the pterygoid [11.1]; (2) a Iacrimal-maxilla suture fused in young adults (reversed in Mesechinus dauuricus) [12.1]; (3) the basioccipital/petrosal suture closed [37.1]; (4) 2 lingual roots of the P4, either fused or not [62]; (5) low trigonids on the lower molars and having the talonid expanded [63.1]; (6) M3 metacone reduced or absent [67.1]. Corbet (1988:119) also listed as derived “the quadrate, low-cusped form of Ml and M2 (with P4 and M3 tending toward that form), the labial styles being reduced to a cingulum; and the subrectangular form of m1 and m2.”

Subfamily HYLOMYINAE Anderson, 1879

GALERICES Pomel, 1848:249. [Type genus: Galerix Pomel, 1848 (fossil). See nomenclatural comment.]

GYMNURINAE Gill, 1872:19. [Type genus: Gymnura Lesson, 1827 (not Gymnura Kuhl, 1823).]

HYLOMIDAE Anderson, 1879:138. [Type genus: Hylomys Müller, 1839.]

ECHINOSORICINAE Cabrera, 1925:57. [Type genus: Echinosorez de Blainville, 1838.]

GALERICINI Butler, 1948:262. [Type genus: Galerix Pomel, 1848. See comment.]

DIAGNOSIS.—(1) rostrum long, greater than 42% of skull length [2.0]; (2) anterior palatine foramina do not include middle palatine foramina [4.0–1]; (3) lacrimal foramen hidden in lateral view by well-developed antorbital flange [8.1]; (4) jugal does not reach posteroventral process of zygoma [10.2]; (5) anterior process of alisphenoid absent [17.0]; (6) suboptic foramen anterior to sphenorbital fissure [20.2]; (7) sphenopalatine foramen anterodorsal or slightly posterodorsal to the palatine transverse torus [21.0]; (8) zygomatic process of squamosal not elevated posteriorly [24.0]; (9) postglenoid foramen posterior to glenoid fossa [25.0]; (10) suprameatal fossa absent [27.0]; (11) nasopharyngeal pocket of basisphenoid absent [30.0]; (12) ectotympanic slender and ring-shaped [32.0]; (13) stapedial foramen posterior to squamosal/alisphenoid suture and posterior to postglenoid foramen [38.0]; (14) epitympanic recess formed by squamosal [39.0]; (15) paroccipital process small [40.0]; (16) occipital condyle emarginated, giving it a slightly lobed appearance [42.0]; (17) condylar foramen of basioccipital anterior to ventral lip of condyle [43.0]; (18) il present [45.0]; (19) i2 subequal to other incisors [47.0]; (20) I2 greater than I3 [48.0]; (21) p3 present [57.0]; (22) M3 hypocone and metacone well developed [66.1, 67.1]; (23) posteroventral keel present on axis [70.1]; (24) metacromion process of scapula elongated, fusiform [71.1]; (25) sacral vertebral not fused into a longitudinal plate [72.1]; (26) greatly elongated posteroventral process of pubis [73.1]; (27) strongly developed lateral flange on anterosuperior margin of tibia [74.1]; (28) pelage not composed of stout spines [77.0].

CONTENT.—Echinosorex de Blainville, 1838; Podogymnura Mearns, 1905; Hylomys Müller, 1839.

DISTRIBUTION.—From northeastern Burma and southern China (Yunnan, Sichuan, Guizhou, Hainan Island) south through Indochina, Indonesia (Sumatra, Borneo, and Java), and the Philippine Islands (Dinagat and Mindanao).

NOMENCLATURAL COMMENT.—The use of the name Galericinae for this taxon is incorrect. Pomel (1848:249) first used the French collective Galerices based on Galerix Pomel, 1848. Unlike other names proposed in the same work, a Latin equivalent was not supplied. According to the International Code of Zoological Nomenclature (1985, Art. ll(f)iii), “A family group name published before 1900 …, but in itself not fully latinized, is available with its original author and date, provided that it has been latinized by later authors and that it has been generally accepted as valid by authors interested in the group concerned and as dating from that first publication as a vernacular name.” However, Galericinae docs not meet these requirements and, in fact, the name was not formally proposed before Butler (1948) as Galericini. Prior to Van Valen (1967), who employed the name Galericinae, the subfamilial name of universal use was Echinosoricinae Cabrera, 1925 (e.g., Simpson, 1945; Findley, 1967; Butler, 1948, 1956), even though this is not the family-group name of priority. Although some workers have followed Van Valen (1967) (e.g., Novacek, 1985, 1986; Corbet, 1988), others (Yates, 1984) have continued to employ Echinosoricinae. Also “galericines” has not been employed as a vernacular name “from that first publication.” Therefore Galericinae is not the family-group name of priority. Surprisingly, when erecting Echinosoricinae for this group, Cabrera (1925) mentioned the family-group name Hylomidae (sic) of Anderson (1879), which is the family-group name of priority.

Genus Echinosorex de Blainville, 1838

Gymnura Lesson, 1827:171 [not Gymnura Kuhl, 1823]. [Type species: Gymnura rafflesii Lesson, 1827 (= Viverra gymnura Raffles, 1822).]

Echinosorex de Blainville, 1838:742. [Type species: Viverra gymnura Raffles, 1822.]

DIAGNOSIS.—(1) palatine foramina small [4.0] and anterior to the maxilla/palatine suture [5.1]; (2) infraorbital canal dorsal or posterodorsal to the P4-M2 region [6.0]; (3) antorbital fossa not present (except weakly in some old adults) [7.1]; (4) posteroventral process of maxillary part of zygoma present, prominent [9.2]; (5) supraorbital processes absent or poorly defined [13.0], bases not formed by anterior processes of parietal [16.0]; (6) ophthalmic foramen joined with or closely adjacent to ethmoid foramen [19.0]; (7) lateral fossa present in palatine anterodorsal to the postpalatal torus [23.1]; (8) coronoid process of dentary broad and rounded [44.1]; (9) 12 larger than 13 [48.0]; (10) C1 significantly larger than adjacent teeth [50.0]; (11) c1 significantly larger than p1 [52.1]; (12) P3 lingual lobe well developed [58.0], P3 normal sized [59.0]; (13) underfur present [81.1].

CONTENT.—Echinosorex gymnura (Raffles, 1822:272) (note: the widespread spelling of the epithet as gymnurus is incorrect; gymnura is a noun in apposition, meaning “nakedtail” and does not change its form to match the gender of the generic name); type locality not given, “Sumatra” implied.

DISTRIBUTION.—Tropical forests of southern Burma and Thailand, Malaya, Sumatra, and Borneo.

DISCUSSION.—The moonrat (or greater gymnure) retains a primitive suite of morphological characters, reminiscent of the common opossums (Didelphis) of the New World. We have been able to identify only a few autapomorphies among the characters that we have analyzed. The lateral fossa of the palatine, which serves for the insertion of the medial pterygoid muscle, is one of the most easily recognized apomorphies of a basically primitive Echinosorex. Butler (1956) pointed out the distinctive eye muscle depression above the optic foramen; however, our examination of other hylomyines reveals this to be a size-related feature. A few dubious qualitative characters might also be considered derived: large body size, Echinosorex being the largest species of insectivore; long tail; possibly aposematic, black-and-white to mostly white pelage (Corbet, 1988); and pre-anal glands that emit a strong ammonia-like scent (Gould, 1978).

In contrast to other insectivores, female Echinosorex are somewhat larger than males (Ralls, 1976). Geographic variation in size has been reported from Thailand, more northerly populations being larger than those adjacent to the equator (Lekagul and McNeeley, 1977).

Genus Podogymnura Mearns, 1905

Podogymnura Mearns, 1905:436. (Type species: Podogymnura truei Meams, 1905.]

DIAGNOSIS.—(1) anterior palatine foramina small [4.0] and anterior to the maxilla/palatine suture [5.1]; (2) infraorbital canal dorsal or posterodorsal to the P4-M1 region [6.0]; (3) antorbital fossa present [7.0]; (4) posteroventral process of maxillary part of zygoma present, small (P. truei) to prominent (P. aureospinula) [9.1–2]; (5) supraorbital processes of frontal absent [13.0], not formed by anterior processes of parietal [16.0]; (6) ophthalmic foramen joined with or closely adjacent to the ethmoid foramen [19.0]; (7) lateral fossa absent from palatine anterodorsal to the postpalatal torus [23.0]; (8) coronoid process of dentary broad and rounded [44.1]; (9) 12 larger than 13 [48.0]; (10) C1 significantly larger than adjacent teeth [50.0]; (11) cl significantly larger than p1 [52.1]; (12) P3 lingual lobe vestigial or absent [58.1], P3 normal sized [59.0]; (13) underfur present [81.1],

CONTENT.—Podogymnura aureospinula Heany and Morgan, 1982:14; type locality: “Plaridel, Albor Municipality, Dinagat Island, Surigao del Norte Province, Republic of the Philippines” [Dinagat Island is at 10°12′N, 125°35′E]. Podogymnura truei Meams, 1905:437; type locality: “Mount Apo at 6,000 feet altitude, southern Mindanao, Philippine Islands” [= 6°59′N, 125°16′E],

DISTRIBUTION.—Dinagat and Mindanao islands, Philippines.

Genus Hylomys Müller, 1839

Hylomys Müller, 1839:50. [Type species: Hylomys suillus Müller, 1839:50]

Neotetracus Trouessart, 1909:389. [Type species: Neotetracus sinensis Trouessart, 1909:389.]

Neohylomys Shaw and Wong, 1959:422. [Type species: Neohylomys hainanensis Shaw and Wong, 1959:422.]

DIAGNOSIS.—(1) anterior palatine foramina elongated posteriorly [4.1] and at the palatine/maxilla suture [5.0]; (2) infraorbital foramen dorsal to the P3–P4 region [6.1]; (3) antorbital fossa present [7.0]; (4) posteroventral process of maxillary part of zygoma absent [9.0]; (5) supraorbital processes of frontal sharp, readily identifiable [13.1], bases formed by long anterior processes of parietals that extend along the supraorbital rim [16.1]; (6) ophthalmic foramen widely separated from the ethmoid foramen [19.1]; (7) lateral fossa absent from palatine anterodorsal to the postpalatal torus [23.1]; (8) coronoid process of dentary narrow and pointed [44.0]; (9) I2 smaller than or subequal to I3 [48.1]; (10) C1 slightly larger than or subequal to adjacent teeth [50.1–2]; (11) c1 approximately equal to, or smaller than, p1 [52.0]; (12) P3 lingual lobe vestigial or absent [58.1], P3 reduced [59.1]; (13) underfur not present [81.0].

CONTENT.—Hylomys hainanensis (Shaw and Wong, 1959:422); type locality: English translation, page 426: “Paisa Hsian, Hainan Island” [= Baisha Xian, which is an administrative unit at 19°13′ 109°26′E]. Hylomys sinensis (Trouessart, 1909:389); type locality: “Ta-tsien-lou, province of Se-tchouen (China Occidental) at an altitude of 2454 meters” [= Kangding, Sichuan Sheng, 30°07′N, 102°02′E]. Hylomys suillus Müller, 1839:50; type locality: “Java.”

FIGURE 11.—Detailed ventral view of skull of Hylomys hainanensis. (Scale = 10 mm; key to abbreviations in Table 2.)

FIGURE 12.—Hylomys hainanensis, A, lateral, B, dorsal, C, occlusal view of lower mandible; D, lateral view of lower mandible. (Scale = 10 mm; key to abbreviations in Table 2).

DISTRIBUTION.—From northeastern Burma, northern Vietnam, and China (Yunnan, Sichuan, Guizhou) (H. sinensis); south through eastern Burma, Thailand, Laos, Vietnam, Malaya, Sumatra, Borneo, and Java (H. suillus); and Hainan Island (H. hainanensis).

COMMENTS.—Hylomys suillus is parapatric with the more northerly H. sinensis; they may have a zone of contact in eastern Burma and northern Thailand (Lekagul and McNeely, 1977) and perhaps Yunnan, although no area of sympatry is yet known.

Subfamily ERINACEINAE Fischer, 1817

ERINACEINAE Fischer. 1817:372. [Type genus: Erinaceus Linnaeus, 1758.]

DIAGNOSIS.—(1) rostrum short, less than 35% of skull length [2.1]; (2) anterior palatine foramina elongated to include middle palatine foramina [4.2]; (3) lacrimal foramen not hidden in lateral view, antorbital flange poorly developed [8.0]; (4) jugal vestigial, not reaching posteroventral process of zygoma [10.1]; (5) anterior process of alisphenoid present [17.1]; (6) suboptic foramen in medial wall of sphenorbital transverse torus [20.1]; (7) sphenopalatine foramen decidedly posterodorsal to palatine transverse torus [21.1]; (8) zygomatic process of squamosal elevated posteriorly [24.1]; (9) postglenoid foramen confluent with glenoid fossa [25.1]; (10) suprameatal fossa moderately to extremely deep [27.1–4]; (11) nasopharyngeal fossa of basisphenoid deep [30.1]; (12) ectotympanic broad [32.1]; (13) stapedial foramen located on squamosal/alisphenoid suture and adjacent to postglenoid foramen [38.1]; (14) epitympanic recess formed by mastoid [39.1]; (15) paroccipital processes robust [40.1]; (16) no emargination of occipital condyle [42.1]; (17) condylar foramen of basioccipital on anteroventral lip of condyle [43.1]; (18) il absent [45.1]; (19) i2 greatly enlarged over other incisors [47.1]; (20) I2 less than or equal to 13 [48.1]; (21) p3 absent [57.2]; (22) M3 hypocone and metacone absent [66.0, 67.0]; (23) posteroventral keel absent from axis [70.0]; (24) metacromion process of scapula deltoid [71.0]; (25) sacral vertebrae fused into a longitudinal plate [72.1]; (26) no greatly elongated posteroventral process of ischium [73.0]; (24) no strongly developed lateral flange on anterosuperior margin of tibia [74.0]; (25) pelage composed of stout spines [77.1–3].

CONTENT.—Atelerix Pomel, 1848; Erinaceus Linnaeus, 1758; Hemiechinus Fitzinger, 1866; Mesechinus Ognev, 1951.

DISTRIBUTION.—Europe, northern Asia south through the Arabian and Indian peninsulas, and into central China; Africa, except for the tropical forest zone.

Genus Hemiechinus Fitzinger, 1866

Ericius Sundevall, 1842:223 [not Ericius Tilesius von Tilenau, 1813]. [Type species: Erinaceus auritus Gmelin, 1770 (= Hemiechinus auritus).]

Hemiechinus Fitzinger, 1866:565. [Type species: Erinaceus plalyolis Sundevall, 1842:232 (= Hemiechinus auritus).}

Paraechinus Trouessart, 1879:242. [Type species: Erinaceus micropus Blyth in Hutton, 1846:170.]

Macroechinus Satunin, 1907:189. [Type species: Erinaceus hypomelas Brandt, 1836:32.]

Erinaceolus Ognev, 1928:132. [Type species: Hemiechinus microtis Laptev, 1925. Nomen inquirenda, see discussion.]

DIAGNOSIS.—(1) rostrum narrow, anterior incisors closely approximated [3.1]; (2) lacrimal/maxilla suture fused and indistinct in young adults [12.1]; (3) posterior palatal shelf with a well-developed spine [22.0]; (4) suprameatal fossa entirely in squamosal [26.0]; (5) suprameatal fossa moderately to extremely well developed [27.2–4]; (6) anterior and posterior borders of suprameatal fossa widely separated [28.0]; (7) parietals relatively higher than frontals [29.0]; (8) basisphenoid inflated [31.1]; (9) squamosal does not participate in bullar roof [33.0]; (10) 13 with two separate roots [49.1]; (11) P2 with 1 root or 2 roots fused [56.1]; (12) long ears [75.1]; (13) hallux reduced [76.1]; (14) pelage spines papillate and grooved [77.3]; (15) ventral pelage soft and densely furred [80.0].

CONTENT.—Hemiechinus auritus (Gmelin, 1770:519); type locality: “in regione Astrachanensi” [Astrakhan’skaya Oblast’, U.S.S.R., 46°21′N 48°03′E]. Hemiechinus collaris (Gray, 1830–1834, plate 8); type locality: “Doab.” Restricted by Wroughton (1910:81) to “between Jumna and Ganges Rivers.” [India]. Subgenus Paraechinus Trouessart, 1879 (see separate account under Paraechinus).

DISTRIBUTION.—Northern Africa eastward through Arabia and the Near and Middle East to India, Soviet Middle Asia, Mongolia, and western China.

DISCUSSION.—Species in the subgenus Paraechinus are treated in a separate account. Hemiechinus auritus and H. collaris, the species not included in the Paraechinus monophyletic group, are treated in this comment Excluding the subgenus Paraechinus, Hemiechinus species have a distribution on Cyprus, and from coastal Libya eastward through the Near and Middle East to northwestern India, and through Soviet Middle Asia to Mongolia and western China.

Corbet (1984, 1988) recognized three species within his Hemiechinus (which he regarded as a distinct genus not including Paraechinus), two quite restricted geographically. The most widely distributed species, H. auritus, occurs from eastern North Africa through the Near and Middle East to Mongolia and western China (Xinjiang), with a southern extension into Pakistan and northwestern India. Populations of eastern Iran, Afghanistan, and Pakistan have been considered by some to be a distinct species, H. megalotis (Lay, 1967; Roberts, 1977). However, Gropp et al. (1969), Hassinger (1973), and Niethamer (1969) provided evidence that the smaller, lighter H. aurilus intergrades with larger, darker populations of H. megalotis in central Iran, and our observations support this. There is, however, an abrupt break in this clinal trend in geographic variation evident along the western side of the Indus River valley. The small, very dark Hemiechinus of the Indus Valley and adjacent northern India also have been considered a subspecies, H. auritus collaris, but no zone of potential contact between the large H. a. megalotis of montane Pakistan and the smaller H. a. collaris in the Indus Valley has been identified (Roberts, 1977), although one may exist The taxa exhibit virtually no overlap in size (Table 8), and also differ in relative ear length (Table 4), length and color pattern of spines, and pelage color of face and throat (Roberts, 1977). No intermediates are known, and given the marked differences between the taxa, they should be retained as separate species, as did Corbet (1988), pending evidence of present or former intergradation. Should this evidence be found, that will constitute evidence of monophyly of Hemiechinus auritus + H. collaris.

TABLE 8.—Cranial measurements of Hemiechinus in Afghanistan and Pakistan.

The taxon microtus is an enigmatic one, originally described by Laptev (1925) as a new species belonging to the genus Hemiechinus. The species is based on only two specimens, both captured alive in “Tashkent city gardens” in Uzbekistan, U.S.S.R. Laptev (1925) based his generic assignment on the absence of a medial spineless tract on the crown, a characteristic of Hemiechinus, but a suite of unusual cranial characters subsequently led Ognev (1928) to erect a monotypic genus, Erinaceolus, to accommodate the species. Bobrinskii et al. (1944:39) next opined that the taxon was based on aberrant individuals because “all the distinctive features…are connected with an anomalous under-development of the hearing apparatus,” and most now regard the name as a synonym of H. auritus.

Subgenus Paraechinus Trouessart, 1879

Paraechinus Trouessart, 1879:242. [Type species: Erinaceus micropus Blyth in Hutton, 1846:170.]

Macroechinus Satunin, 1907:189. [Type species: Erinaceus hypomelas Brandt, 1836.]

DIAGNOSIS.—(1) alisphenoid inflated [11.2] (not so in other Hemiechinus); (2) suprameatal fossa very deep [27.3] (less so in other Hemiechinus); (3) ventral process of petrosal inflated [36.1]; (4) spineless tract on crown wide [79.2] (tract absent in other Hemiechinus).

CONTENT.—Hemiechinus aethiopicus (Ehrenberg, 1833, Dec 2, leaf k recto (footnote)); type locality: “In desertis dongolanis habitat.” [approximately at 19°22′N, 30°45′E]. Hemiechinus hypomelas (Brandt, 1836:32); type locality: “pays des Turcomans” [the “country of the Turkmens” is roughly equivalent to the Turkmen, Uzbek, Kazak, and Karakalpak regions of the U.S.S.R.]. Hemiechinus micropus (Blyth in Hutton, 1846:170); type locality: “Bhawulpore” [= Bahawalpur, Pakistan, 29°23′N, 71°39′E]. Hemiechinus nudiventris (Horsfield, 1851:136); type locality: “Madras” [“Madras” could refer to the city at 13°05′N, 80°17′E or the older name for the Tamil Nadu province].

DISTRIBUTION.—Northern Africa, south to Mauritania and northern Somalia, and east throughout the Arabian Peninsula and Middle Asia to western and southern India.

DISCUSSION.—The subgenus Paraechinus shows considerably more size variation when compared to other Hemiechinus (Table 8). All species have a well-defined basisphenoid fossa and a greatly enlarged hypotympanic sinus. This sinus is extended into the alisphenoid and pterygoids such that they form a significant part of the anterior part of the sinus. Some characters that have been used to separate species (i.e., maxilla-nasal contact, palatine transverse torus straight or curved) we found too variable within taxa to be of diagnostic use. This subgenus also shows the greatest development of the auditory region of any insectivore. The hypotympanic chamber has expanded laterally, dorsally, and anteriorly to form an extremely large sinus. Within this group, H. aethiopicus and H. micropus show the most extreme development.

Hemiechinus (Paraechinus) aethiopicus (sensu lato) ranges throughout North Africa and eastward through Sinai and Arabia to Iraq, but only as far as the Tigris-Euphrates region. It is usually light in color, with light-tipped spines, a robust skull with broad maxilla, a relatively high, more dome-shaped braincase, very inflated auditory region, and large teeth (Table 6), but P3 is reduced to a single cusp, or absent. Osborn and Helmy (1980), following Setzer (1957), recognized within the former H. (P.) aethiopicus three species (H. (P.) aethiopicus, H. (P.) deserti, and H. (P.) dorsalis). These three putative species are mutually allopatric. In the absence of critical data from zones of potential contact between these taxa, and in the light of the considerable variation evident in some of the characters evinced by these taxa, we consider the three as subspecies of H. aethiopicus.

Hemiechinus a. deserti is widespread in North Africa, from Mauritania and Morocco in the west to the Nile in the east. Osborn and Helmy (1980) distinguished it from others on the basis of its pale spine tips, lesser parapterygoid inflation, a small medial cusp on p4, straight postpalatal bridge, and the maxillae not contacting the nasals. While this diagnosis holds true of specimens from Egypt that we have examined, those from farmer west in Libya (Hufnagel, 1972), Morocco, Niger, and Mauritania are more variable in color. A series of 22 from the western end of the taxon’s distribution included 17 that were typically pale, but five possessed dark spines, including the tips. Moreover, the relationship of maxillae to nasals (Table 5), degree of parapterygoid inflation, and size of medial cusp on p4 are also individually variable in these western populations.

Hemiechinus a. aethiopicus occurs east of the Nile and south of the delta region in the Eastern Desert of Egypt, extending southward to northern Ethiopia and Somalia. Its westward extension in Sudan, and possible contact with P. a. deserti, which is known from Niger, is unknown. Osborn and Helmy (1980) believed it distinctive on the basis of its dark spine tips, relatively greater parapterygoid inflation, the absence of a medial cusp on p4, straight postpalatal bridge, and the maxillae contacting the nasals. In the specimens that we examined (Appendix 1), these characters seem to hold, except for the degree of inflation of the parapterygoids, which was variable, and not much different from the condition seen in H. a. deserti.

Hemiechinus a. dorsalis occurs from the Sinai on the west throughout the Arabian Peninsula, and north and eastward to the Tigris-Euphrates region of Iraq. Contact with H. a. aethiopicus is presumed to have been in the Suez region. Osbom and Helmy (1980) described it as the darkest of the three taxa, with dorsal spines entirely dark, relatively moderate parapterygoid inflation, the medial cusp on p4 vestigial, the maxillae contacting the nasals, and a slightly V-shaped postpalatal bridge. However, coloration is again quite variable, with some specimens being considerably lighter than usual (e.g., USNM 321572). We found the variation in the pterygoid inflation to be large (to be sure, correlated with the great amount of variation typical of this subgenus) and very subtle among the allopatric populations. There was considerable variation in the shape of the postpalatal bridge and nasal-maxilla contact (Table 5).

Hemiechinus (Paraechinus) micropus is found in the lower Indus Valley of Pakistan (Roberts, 1977) and adjacent India south to Gujarat. It is a small, pale-colored hedgehog with relatively short ears (Table 2), short light-tipped spines, and the “same mask pattern [that] occurs in P. aethiopicus…and may be conspecific” (Roberts, 1977:20). The skull resembles that of H. aethiopicus in being robust with a relatively short rostrum, with broad maxillae, broad high braincase, and relatively large teeth (Table 6), but a reduced P3. Corbet (1988) distinguishes between H. aethiopicus and H. micropus on the relative inflation of the mastoid. We could discern no difference in the relative inflation of the mastoid per se; however, the suprameatal fossa (with mastoid and squamosal components) is greatly inflated in these taxa, with H. hypomelas with the smallest and H. aethiopicus with the greatest. In H. micropus and H. aethiopicus, this cavity is further enlarged to extend posterodorsal to the glenoid fossa. Hemiechinus (P.) micropus probably represents a peripheral isolate of H. (P.) aethiopicus, whose ranges are now separated by a gap occupied by H. (P.) hypomelas and H. auritus.

Biswas and Ghose (1970) proposed that specimens from Rajasthan, Madhya Pradesh, and Gujarat represent a second species, H. intermedius, which is, in part, sympatric with H. micropus. They distinguished H. intermedius on the basis of smaller size and different color pattern, especially on the face. However, facial pattern is variable (Roberts, 1977) and the sample of cranial measurements is small (N=4); we have not examined this form, and provisionally consider it a synonym of H. micropus, as did Corbet (1988).

Neither have we examined specimens of H. (P.) nudiventris, which Biswas and Ghose (1970) also considered a distinct species (but which Corbet, 1978, 1988, considered to be a subspecies of H. (P.) micropus). It is an isolated population, restricted to the states of Madras (= Tamil Nadu) and Travancore (= Kerala) in southeastern India. This taxon differs from H. (P.) micropus in possessing a narrower cranium, incomplete zygomatic arch, and vestigial P3 (which Biswas and Ghose referred to as P2) in the two skulls that they examined. It also differs in size and shape of ear, tail length, and sole of the forefoot. The vestigial P3 is clearly a derived condition. As described by Biswas and Ghose (1970) the incomplete zygomatic arch is the result of the absence of the “malar” (= jugal) bone, and separation of the zygomatic processes of the maxilla and squamosal. This condition, if not an artifact of preparation, is unique to the family, and clearly an autapomorphy; H. (P.) nudiventris is therefore provisionally considered a distinct species, possibly derived from or allied with H. (P.) micropus.

Hemiechinus (Paraechinus) hypomelas is found from the Transcaspian and Aral regions southward through Iran and eastward to the Indus River in Pakistan. It is also reported from the Punjab in northwestern India (Roberts, 1977; Corbet, 1978). It is the only species of hedgehog that exhibits color dimorphism. Most populations are dark with dark-tipped spines, but populations containing nearly white individuals occur in the northern and central part of its range. The skull is relatively light, with narrow maxilla and a flat braincase, and small teeth, but the P3 has three distinct cusps. In Pakistan, two populations of dark-colored H. (P.) hypomelas occur that differ greatly in size. Hemiechinus h. hypomelas is large and found in the foothills of western Pakistan, whereas H. h. blanfordi (= jerdoni, see below) is small with relatively small ears (Tables 4 and 8), and short spines (Roberts, 1977), and is found in the Indus Valley. The ranges of size overlap only slightly (Table 8). Thus, these taxa parallel in size and distribution Hemiechinus auritus megabits and H. collaris occurring in the same geographic area (see above). As in that case, no zone of potential contact is known.

G.B. Corbet (in litt.) has informed us that H. h. blanfordi Anderson (1878) “is a Hemiechinus collaris as is clear from the original description and as pointed out by Agrawal (1973)….” Thus, the name jerdoni Anderson (1878) must be employed instead for the small dark subspecies of H. (P.) hypomelas that inhabits the Indus Valley. Externally, H. (P.) hypomelas jerdoni and H. collaris are very similar, and in the zone of sympatry may be difficult to distinguish. Nevertheless, H. (P.) hypomelas possesses a well-defined medial spineless tract on the crown, rugose rather than smooth spines, and a reduced hallux, all characters that arc not present in H. collaris.

Genus Mesechinus Ognev, 1951

Mesechinus Ognev, 1951:8. [Type species: Erinaceus dauuricus Sundevall, 1842:237.]

DIAGNOSIS.—(1) rostrum broad, anterior incisors not closely approximating [3.0]; (2) lacrimal/maxilla suture not fused in young adults [12.0]; (3) posterior palatal shelf with a well-developed spine [22.0]; (4) suprameatal fossa entirely in squamosal [26.0]; (5) supramcatal fossa shallow [27.1]; (6) anterior and posterior borders of suprameatal fossa narrowly separated [28.1]; (7) parietals relatively higher than frontals [29.0]; (8) basisphenoid not inflated [31.0]; (9) squamosal forming a major part of bullar roof [33.1]; (10) I3 with 2 separate roots [49.1]; (11) P2 with 1 or 2 roots fused [56.1]; (12) ears short [75.0]; (13) hallux reduced [76.1]; (14) pelage spines papillate, not grooved [77.2]; (15) ventral pelage coarse [80.1].

CONTENT.—Mesechinus dauuricus (Sundevall, 1842:237); type locality: “Dauuria” [U.S.S.R., Transbaikalia, Dauryia, 49°57′N, 116°55′E]. Mesechinus hughi (Thomas, 1909:966); type locality: “Paochi, Shen-si” [= Baoji, Shaanxi Sheng, China, 34°23′N 107°09′E].

DISTRIBUTION.—Northeastern and central Mongolia and limited adjacent areas in China and the U.S.S.R. (M. dauuricus); in Shanxi and Shaanxi of north central China (M. hughi).

DISCUSSION.—Bobrinskii et al. (1965) considered M. dauuricus to be a subspecies of Erinaceus europaeus, although most authors had placed it in the genus Hemiechinus (Stroganov, 1957; Corbet, 1978). Subsequently, Zaitsev (1982) and Pavlinov and Rossolimo (1987) employed the subgenus Mesechinus Ognev, 1951, within Erinaceus, for this species.

South and west of the range of M. dauuricus, in central China (Shanxi and Shaanxi), another hedgehog occurs, originally named Erinaceus hughi (Thomas, 1909). Corbet (1978) reallocated it to Hemiechinus, and later (1984) elevated it to full species status and also included Hemiechinus sylvaticus (Ma, 1964) as a junior synonym. The holotype of hughi is a skin only, and the taxon is apparently known from only two localities. We have examined the holotypcs of both named forms, and concur with Corbet (1978) that hughi (including sylvaticus) probably represents a distinct species. It shares with M. dauuricus absence of a median spineless tract, a possible synapomorphy with Hemiechinus (not including Paraechinus), or (following Corbet, 1988) a character plesiomorphic for erinaceines.

The form miodon of northern Shaanxi has been assigned in the past to M. dauuricus but is separated from Mongolian populations of M. dauuricus by nearly 1600 km of apparently unsuitable semidesert and desert. It is much closer geographically to M. hughi, and its relationships should be re-examined.

Mesechinus is distinctive in many features, most notably the unique shape of the suprameatal fossa. Whereas in most crinaceines, the lateral borders of this fossa are somewhat C-shaped, in Mesechinus, the anterior and posterior rim are nearly parallel, giving the fossa a more angular or U-shaped appearance. The basisphenoid inflation is intermediate between the condition more typical of Hemiechinus and that found in Atelerix and Erinaceus (Figure 4). The development of the lingual lobe on P3 was individually variable among the specimens we examined.

Genus Erinaceus Linnaeus, 1758

Erinaceus Linnaeus, 1758:52. [Type species: Erinaceus europaeus Linnaeus, 1758.]

DIAGNOSIS.—(1) rostrum broad, anterior incisors not closely approximating [3.0]; (2) lacrimal/maxilla suture fused and indistinct in young adults [12.1]; (3) posterior palatal shelf with a well-developed spine [22.0]; (4) suprameatal fossa entirely in squamosal [26.0]; (5) suprameatal fossa shallow [27.1]; (6) anterior and posterior borders of suprameatal fossa widely separated [28.0]; (7) frontals relatively higher than parietals [29.1]; (8) basisphenoid not inflated [31.0]; (9) squamosal not participating in bullar roof [33.0]; (10) 12 with 2 roots fused [49.2]; (11) P2 with 1 root or 2 roots fused [56.1]; (12) ears short [75.0]; (13) hallux normal [76.0]; (14) pelage spines smooth [77.1]; (15) ventral pelage coarse [80.0].

CONTENT.—Erinaceus europaeus Linnaeus, 1758:52; type locality: “Europa.” Erinaceus concolor Martin, 1838:103]; type locality: “apud Trebizond” [= Trabzon, Turkey, 40°00′N, 39°43′E]. Erinaceus amurensis Schrenk, 1859:100; type locality: “In der Nähe der Stadt Aigun, im mandshurischen Dorfe Guissoja am Amur ….”

DISTRIBUTION.—Throughout the western Palearctic, from Ireland and Spain to Turkey, the western shore of the Caspian Sea, and central Siberia (80° E long.) (E. europaeus, E. concolor). A separate area is occupied by E. amurensis in the Korean peninsula and adjacent Manchuria, and from the vicinity of Beijing south to the Yangtze River valley, and westward to the vicinity of western Hubei province. The status of specimens reported from farther west is unclear (see Corbet, 1988).

DISCUSSION.—Most authors now agree that two sibling species of hedgehogs occur in Europe, the western and northern E. europaeus and the eastern and southern E. concolor (Corbet, 1984, 1988). Their distribution is parapatric to narrowly sympatric in Poland, Czechoslovakia, and Austria (Ruprecht, 1973; Kratochvil, 1975; Puček and Raczynski, 1983; Corbet, 1988) and perhaps elsewhere, where they are distinguishable on the basis of ventral coloration, and maxillary and mandibular structure (Puček, 1981; Zaitsev, 1984; Corbet, 1988). Corbet (1988) separated E. concolor and E. europaeus on two features of the rostrum, both involving the posterior extension of the maxilla. Although there appears to be a general trend to support E. concolor having a longer naso/maxilla suture than E. europaeus, we found this feature variable (Table 5). However, we found no exceptions to Corbet’s notice of the maxilla extending farther posterior in E. concolor than in E. europaeus.

In the Far East, other populations of Erinaceus exist widely disjunct from the two European species, ranging from the Amur River drainage of southern Siberia on the north, south throughout the Korean Peninsula and Manchuria, to the northeastern provinces of China. These were allocated to E. europaeus in the past, before E. concolor was recognized as distinct. More recently, Far Eastern hedgehogs have been elevated to specific status as E. amurensis (Corbet, 1984; Zaitsev, 1984), including the possibly allopatric southern populations (E. dealbatus). In E. amurensis and E. concolor the maxilla/premaxilla suture is nearly perpendicular to the nasals, whereas in E. europaeus, the sutures meet at an acute angle. Additionally, E. amurensis shares with E. concolor the tendency for the posterior extension of the maxilla beyond the lacrimal foramen (Puek, 1981; Zaitsev, 1984). Whether these features are due to synapomorphy or homoplasy, we do not know, although Corbet (1988) interpreted these as synapomorphies. The Far Eastern species also differs from the two European taxa in having a shorter ear, both absolutely and relatively (Table 4), a V-shaped basisphenoid fossa (Corbet, 1988), and a consistently light venter, and the face is also usually light, though these last two characters show some variation. All three species possess a dark snout.

Genus Atelerix Pomel, 1848

Atelerix Pomel, 1848:251. [Type species: Erinaceus albiventris Wagner, 1841, by subsequent designation of Thomas (1918).]

Peroechinus Fitzinger, 1866:565. (Type species: Erinaceus pruneri Wagner, 1841 (= Atelerix albiventris).]

Aethechinus Thomas, 1918:194. [Type species: Erinaceus algirus Lereboullet in Duvemoy and Lereboullet, 1842.)

DIAGNOSIS.—(1) rostrum broad, anterior incisors not closely approximating [3.0]; (2) lacrimal/maxilla suture fused and indistinct in young adults [12.1]; (3) posterior palatal shelf without a well-developed spine [22.1]; (4) suprameatal fossa with mastoid contribution greater than or subequal to squamosal contribution [26.1]; (5) suprameatal fossa shallow [27.1]; (6) anterior and posterior borders of suprameatal fossa widely separated [28.0]; (7) frontals relatively higher than parietals [29.1]; (8) basisphenoid not inflated [31.0]; (9) squamosal not participating in bullar roof [33.0]; (10) I3 with 2 roots separate [49.1]; (11) P2 with 2 roots [56.0]; (12) ears short [75.0]; (13) hallux reduced or absent [76.1]; (14) pelage spines smooth [77.1]; (15) ventral pelage coarse [80.1].

CONTENT.—Atelerix albiventris (Wagner, 1841:22); type locality: unknown; Wagner stated that the holotype arrived with a shipment of specimens from India. Anderson and de Winton (1902) believed it to be from “Senegambia.” Atelerix algirus (Lereboullet in Duvemoy and Lereboullet, 1842:4); type locality: “provient d’Oran” [Algeria, currently the province Wilaya d’Oran]. Atelerix frontalis (Smith, 1831); type locality: “Cape” [Province, South Africa]. Atelerix sclateri (Anderson, 1895:415); type locality: “Taf in Central Somaliland.”

DISTRIBUTION.—Northwestern Africa to eastern Libya; Senegal to Somalia; southern Angola and Zimbabwe south to South Africa; in East Africa south to Malawi. Also on Canary and certain Mediterranean islands and Mediterranean coast of Spain and France, where probably introduced.

DISCUSSION.—Aethechinus has usually been considered a synonym of Erinaceus (Corbet, 1978, 1984; Smithers, 1983), but recently Robbins and Setzer (1985) advanced arguments supporting its distinctness and included two species: Aethechinus algirus from northern Morocco to Libya and (probably introduced) southern Spain and France as well as the Balearic and Canary Islands and Malta; and A. frontalis, from southern Africa. Robbins and Setzer (1985) concluded, on the basis of a morphometric analysis, that A. frontalis and A. algirus were closest to Erinaceus, while Atelerix albiventris was closest to Paraechinus and Hemiechinus; they recommended that Aethechinus and Atelerix be recognized as separate genera. As previously noted, our phylogenetic analysis did not produce results concordant with their phenetic analysis. The phenetic similarity exhibited by Erinaceus and Aethechinus (sensu Robbins and Setzer) appears to be due to similarity in size; as they noted, Erinaceus spp., Aethechinus frontalis, and Aethechinus algirus are larger, while Atelerix albiventris is the smallest species of spiny hedgehog. Corbet (1988) supplied a third alternative, placing A. algirus and A. frontalis within Atelerix, along with A. albiventris and A. sclateri.

Gropp and Natarajan (1972:267) found that A. algirus shared with Erinaceus the presence of “sizable blocks of distally located heterochromatin,” two in the former and three in the latter, which are not present in Hemiechinus (species not noted). They interpreted this as indicating close relationship between A. algirus and Erinaceus. Unfortunately, they did not study A. albiventris.

The widespread African hedgehog Atelerix albiventris occurs south of the Sahara and north of the forest zone from Senegal eastward to Somalia, and southward in East Africa to Tanzania (Corbet, 1974), Malawi, and to the Zambezi River (central Mozambique) (Corbet, 1988). Its distribution is allopatric with the other species of African hedgehogs, except for a narrow zone of potential contact in Sudan and perhaps Ethiopia. However, the poorly known taxon A. sclateri occurs in northern Somalia, and may be parapatric or sympatic with A. albiventris. In this study, we were only able to examine one young adult male and, therefore, could not take into account interspecific variation. The characters of A. sclateri include the primitive well-developed (though small) hallux (Corbet, 1974, 1988) (also in A. algirus and A. frontalis) and the derived reduced P3 (also in A. albiventris). The shape of the parietal/occipital suture (Corbet, 1974) is too variable in other taxa to be of diagnostic use. The relationship of nasal to maxilla (Corbet, 1974) was individually variable in most specimens of erinaceines examined (Table 5); however, as pointed out by Corbet (1988), our specimen of A. sclateri did not have a maxilla-nasal contact. Our specimen of A. sclateri differed from other Atelerix in other features as well. P3 in A. sclateri is an extremely small, peg-like tooth, and is considerably less than one-half the size of P2. In A. frontalis, although P3 is reduced, it is closer in size to P2. Atelerix sclateri also has a uniquely shaped nasal. As a reflection of the posterior growth of the rostral process of the premaxilla, the nasals are widest posteriorly at the point that the premaxilla and frontals meet. In other Atelerix the nasals are nearly parallel sided.

Ventral coloration is dark in A. frontalis, light in A. albiventris, and variable in A. algirus and A. sclateri; dark ventral pelage is probably primitive, because it is found widely in other hedgehog genera. The few available specimens of A. sclateri (Corbet, 1974, 1988; Kingdon, 1974) suggest that it is the sister taxon of A. albiventris, which retains a few plesiomorphies. Additional specimens may show, however, that it intergrades with typical A. albiventris in southern Somalia and northern Kenya.

As in the A. albiventris-A. sclateri clade, A. frontalis can be separated from A. algirus on the relative sizes of P2 and P3 and the morphology of m3. P2 in A. frontalis is a robust tooth, being larger than either adjacent tooth, whereas in A. algirus, P2 is somewhat smaller and similar in morphology to C1. Although P3 is reduced, in both species there is a well-developed lingual lobe.

Corbet (1988) distinguished among Atelerix on the basis of the naso/maxilla suture. We found the individual variation to mask his implied morphological gaps (Table 5). However, we found no exceptions to his characterization of A. frontalis as having a long common suture and A. sclateri with no common suture.

Appendix 1

List of Specimens Examined

AMNH = American Museum of Natural History; ANSP = Academy of Natural Sciences of Philadelphia; BEIJ = Beijing, Institute of Zoology; BMNH = British Museum (Natural History); DMNH = Delaware Museum of Natural History; FMNH = Field Museum of Natural History; HMCZ = Harvard University, Museum of Comparative Zoology; KU = University of Kansas, Museum of Natural History; MGU = Moscow State University, Zoological Museum; UPLB = University of the Philippines at Los Baños, Museum of Natural History; USNM = former collections of the United States National Museum, now deposited at the National Museum of Natural History, Smithsonian Institution. Arranged alphabetically by taxon and locality, and grouped by sex within each locality.

Atelerix albiventris

USNM

421118

?

Benin: Borgou Region

USNM

421120

?

Benin: Borgou Region

USNM

421136

Benin: Borgou Region

USNM

421138

Benin: Borgou Region

USNM

421127

Benin: Borgou Region

USNM

421132

Benin: Borgou Region

USNM

439833

Ghana: Eastern Region

USNM

424630

Ghana: Northern Region

USNM

439834

Ghana: Northern Region

USNM

424631

Ghana: Northern Region

USNM

a470571

Ghana: Northern Region

USNM

161699

Kenya: Kapita Plains

USNM

181442

Kenya: Lime Springs

KU

41112

?

Kenya: Nairobi Area

USNM

164022

Kenya: Ulucania Hills

USNM

402202

Nigeria: Northern Region

USNM

402203

Nigeria: Northern Region

USNM

483167

Nigeria

Alelerix algirus

USNM

302065

Libya: Bin Jawwad, Baladiyat

USNM

476060

Morocco: Agadir, Province d'

USNM

476058

Morocco: Agadir, Province d'

USNM

476055

Morocco: Fes, Province de

USNM

476052

Morocco: Ksar-es-souk, Province de

USNM

476053

Morocco: Ksar-es-souk, Province de

USNM

470573

Morocco: Ouarzazate, Province d'

USNM

470572

Morocco: Ouarzazate, Province d'

USNM

470574

Morocco: Ouarzazate, Province d'

USNM

470575

Morocco: Ouarzazate, Province d'

USNM

b470571

Morocco: Ouarzazate, Province d'

USNM

476050

Morocco: Oujda, Province d'

USNM

476057

Morocco: Oujda, Province d'

USNM

476062

Morocco: Tetouan, Province de

Atelerix frontalis

USNM

260742

Botswana: Gaberones

USNM

380487

South Africa: Orange Free State

USNM

468211

South Africa: Transvaal

USNM

267653

South Africa: Transvaal

Atelerix sclateri

USNM

63220

Somalia: Woqooyi Galbeed, Gobolka

Echinosorex gymnura

USNM

145585

?

Indonesia: Kalimantan

KU

92668

?

Malaysia: Sabah

KU

92669

Malaysia: Sabah

USNM

145584

Malaysia: Sabah

USNM

487887

?

Malaysia: West

USNM

357885

Malaysia: West

USNM

357887

Malaysia: West

USNM

487886

Malaysia: West

USNM

487889

Malaysia: West

USNM

487892

Malaysia: West

USNM

487898

Malaysia: West

USNM

487899

Malaysia: West

USNM

487900

Malaysia: West

USNM

487902

Malaysia: West

USNM

487903

Malaysia: West

USNM

487894

Malaysia: West

USNM

487897

Malaysia: West

USNM

487901

Malaysia: West

USNM

83499

?

Thailand: Trang, Changwat

USNM

86785

?

Thailand: Trang, Changwat

Erinaceus amurensis

AMNH

57219

China: Anhui Sheng

BEIJ

60000

China: Beijing Shi

BMNH

61.6.2.5

?

China: Beijing Shi

BEIJ

15856

China: Jilin Sheng Dixing

BEIJ

12741

China: Jilin Sheng Dixing (?)

BEIJ

09341

China: Jilin Sheng Dixing

AMNH

56095

China: Hebei Sheng

USNM

176251

China: Hebei Sheng

USNM

199681

China: Hebei Sheng

BEIJ

13305

China: Heilongjiang Sheng Dixing

USNM

270542

China: Heilongjiang Sheng Dixing

BEIJ

15854

China: Henan Sheng Dixing

HMCZ

7132

China: Hubei Sheng

AMNH

57218

China: Hunan Sheng

USNM

239591

China: Hunan Sheng

USNM

239592

China: Hunan Sheng

USNM

239590

China: Hunan Sheng

ANSP

20447

China: Jiangsu Sheng

ANSP

20446

China: Jiangsu Sheng

USNM

174605

?

China: Jiangsu Sheng

USNM

240325

?

China: Jiangsu Sheng

USNM

252158

?

China: Jiangsu Sheng

BMNH

10.5.1.2

China: Jilin Sheng

USNM

197779

China: Jilin Sheng

USNM

270541

China: Liaoning Sheng

ANSP

16684

China: Shandong Sheng

BMNH

8.2.8.2

China: Shandong Sheng

BEIJ

05120

China: Shandong Sheng

BEIJ

05122

China: Shandong Sheng

BEIJ

05123

China: Shandong Sheng

BEIJ

05124

China: Shandong Sheng

ANSP

16679

China: Shandong Sheng

ANSP

16680

China: Shandong Sheng

ANSP

16681

China: Shandong Sheng

ANSP

16682

China: Shandong Sheng

BMNH

8.2.8.1

China: Shandong Sheng

BEIJ

05121

China: Shandong Sheng

BEIJ

05125

China: Shandong Sheng

ANSP

16683

?

China: Shandong Sheng

BMNH

74.1.24.23

7

China: Shandong Sheng

HMCZ

25884

?

China: Shandong Sheng

HMCZ

25885

?

China: Shandong Sheng

BEIJ

0001

China: Shanghai Shi

BEIJ

356

China: Shanghai Shi

BEIJ

0002

China: Shanghai Shi

BEIJ

354

China: Shanghai Shi

USNM

239770

?

China: Zhejiang Sheng

BMNH

22.10.6.2

?

Korea

HMCZ

37692

?

Korea

USNM

302902

?

Korea

USNM

302903

?

Korea

FMNH

48372

Korea

HMCZ

37691

Korea

BMNH

22.10.6.1

Korea

FMNH

48371

Korea

HMCZ

37690

Korea

AMNH

18355

U.S.S.R.: R.S.F.S.R., Primorsk. krai

BMNH

1938.3.30.6

U.S.S.R.: R.S.F.S.R., Primorsk. krai

HMCZ

24731

U.S.S.R.: R.S.F.S.R., Primorsk. krai

Erinaceus concolor

FMNH

96338

Iran: Azarbaijan-e Gharbi, Ostan-e

FMNH

96339

Iran: Azarbaijan-e Gharbi, Ostan-e

FMNH

96340

Iran: Azarbaijan-e Gharbi, Ostan-e

FMNH

96341

Iran: Azarbaijan-e Gharbi, Ostan-e

FMNH

96344

Iran: Azarbaijan-e Gharbi, Ostan-e

FMNH

96345

Iran: Azarbaijan-e Gharbi, Ostan-e

FMNH

110955

Iran: Azarbaijan-e Gharbi, Ostan-e

FMNH

92893

Iran: Kerman, Ostan-e

FMNH

92894

Iran: Kerman, Ostan-e

USNM

369533

Iran: Khuzestan, Ostan-e

FMNH

84448

Iraq

USNM

37464

?

Lebanon

FMNH

94242

Lebanon

FMNH

94244

Lebanon

HMCZ

8593

?

Palestine

HMCZ

12436

?

Palestine

HMCZ

12602

?

Palestine

HMCZ

12603

?

Palestine

HMCZ

17333

?

Palestine

HMCZ

17331

?

Palestine

HMCZ

17332

?

Palestine

HMCZ

17334

?

Palestine

FMNH

74343

Turkey: Ankara

FMNH

83350

Turkey: Ankara

FMNH

82113

Turkey: Bitlis

FMNH

82121

Turkey: Bitlis

FMNH

82122

Turkey: Bitlis

FMNH

82124

Turkey: Bitlis

FMNH

82127

Turkey: Bitlis

FMNH

82110

Turkey: Elazig Ili

FMNH

82109

Turkey: Elazig Ili

FMNH

82111

Turkey: Elazig Ili

FMNH

82112

Turkey: Elazig Ili

HMCZ

51529

Turkey: Elazig Ili

FMNH

74338

Turkey: Hatay Hi

FMNH

74341

Turkey: Hatay Ili

FMNH

74340

Turkey: Hatay Ili

FMNH

74342

Turkey: Hatay Ili

FMNH

82115

Turkey: Hatay Hi

USNM

327188

Turkey: Icel

USNM

327187

Turkey: Istanbul, Ili

USNM

327190

Turkey: Istanbul, Ili

USNM

327189

Turkey: Kayseri

FMNH

82126

Turkey: Urfa

Erinaceus europaeus

USNM

174830

Denmark

USNM

174831

Denmark

USNM

251763

?

Germany

USNM

112336

Germany

USNM

112338

Germany

USNM

153413

Italy: Roma, Provincia di

USNM

154161

?

Spain: Burgos, Provincia de

USNM

154162

Spain: Burgos, Provincia de

USNM

153417

Spain: Burgos, Provincia de

KU

11420

?

Spain: Madrid, Provincia de

USNM

153414

Spain: Oviedo, Provincia de

USNM

153415

Spain: Oviedo, Provincia de

USNM

186554

Sweden: “Uppsala”

USNM

186555

Sweden: “Uppsala”

USNM

85086

Sweden: “Uppsala”

USNM

85090

Sweden: “Uppsala”

USNM

85092

Sweden: “Uppsala”

USNM

847740

Switzerland: Appenfell, Canton

USNM

115198

Switzerland: Saint Gallen, Canton

USNM

105881

?

Switzerland: Saint Gallen, Canton

USNM

84739

?

Switzerland: Saint Gallen, Canton

USNM

115196

?

Switzerland: Saint Gallen, Canton

USNM

115197

?

Switzerland: Saint Gallen, Canton

USNM

115199

Switzerland: Tessin (Ticino), Canton

USNM

795

?

United Kingdom: England

USNM

153410

United Kingdom: England

USNM

174660

United Kingdom: England

USNM

153409

United Kingdom: Wales

USNM

251643

?

Unknown

HMCZ

51529

Turkey

Hemiechinus aethiopicus

USNM

311732

Egypt

USNM

311737

?

Egypt: Sahra' al Gharbiyah, Muhafazat

USNM

311738

?

Egypt: Sahra' al Gharbiyah, Muhafazat

USNM

311731

Egypt: Sharqiyah, as Sahra' ash

USNM

325906

①

Egypt: Sina, Muhafazat

USNM

325908

Egypt: Sina, Muhafazat

USNM

311739

Egypt: Sina, Muhafazat

USNM

325907

Egypt: Sina, Muhafazat

USNM

311740

Egypt: Sina, Muhafazat

FMNH

123048

Egypt: Sina, Muhafazat

BMNH

21.12.3.1

Iran: Tanb Island

USNM

384832

Mauritania: Adrar, Cercle de 1'

USNM

410873

?

Mauritania: Assaba, Cercle de 1'

USNM

410873

Mauritania: Assaba, Cercle de 1'

USNM

410872

Mauritania: Assaba, Cercle de 1'

USNM

470569

Morocco: Ouarzazate, Province de

USNM

470566

Morocco: Ouarzazate, Province de

USNM

470567

Morocco: Ouarzazate, Province de

USNM

470568

Morocco: Ouarzazate, Province de

USNM

482512

Niger

BMNH

52.1468

Oman

AMNH

166942

?

Saudi Arabia

BMNH

1934.8.4.4

?

Saudi Arabia

BMNH

48.378

Saudi Arabia: Jawf, Mintaqat al

BMNH

10.6.30.3

?

Yemen (Aden)

BMNH

10.6.30.4

Yemen (Aden)

BMNH

10.6.30.5

Yemen (Aden)

USNM

321572

Yemen (Aden)

BMNH

54.1020

Yemen (Aden)

BMNH

10.6.30.1

Yemen (Aden)

BMNH

10.6.30.2

Yemen (Aden)

Hemiechinus auritus

BMNH

86.10.15.6

Afghanistan: Badghis, Velayat-e

FMNH

102001

Afghanistan: Baghlan, Velayat-e

FMNH

102008

Afghanistan: Baghlan, Velayat-e

FMNH

102011

Afghanistan: Baghlan, Velayat-e

FMNH

102012

Afghanistan: Baghlan, Velayat-e

FMNH

102013

Afghanistan: Baghlan, Velayat-e

FMNH

48483

Afghanistan: Baghlan, Velayat-e

FMNH

102003

Afghanistan: Baghlan, Vclayat-e

FMNH

102004

Afghanistan: Baghlan, Velayat-e

FMNH

102005

Afghanistan: Baghlan, Velayat-e

FMNH

102007

Afghanistan: Baghlan, Velayat-e

FMNH

102009

Afghanistan: Baghlan, Velayat-e

FMNH

102014

Afghanistan: Baghlan, Velayat-e

FMNH

102015

Afghanistan: Faryab, Vclayat-e

FMNH

102016

Afghanistan: Faryab, Velayat-e

FMNH

102017

Afghanistan: Herat, Velayat-e

FMNH

102021

Afghanistan: Herat, Velayat-e

FMNH

102020

Afghanistan: Herat, Velayat-e

BMNH

47.360

Afghanistan: Kabol, Velayat-e

FMNH

102022

Afghanistan: Kabol, Velayat-e

FMNH

102026

Afghanistan: Kabol, Velayat-e

FMNH

102027

Afghanistan: Kabol, Velayat-e

FMNH

102029

Afghanistan: Kabol, Velayat-e

BMNH

47.359

Afghanistan: Kabol, Velayat-e

FMNH

102023

Afghanistan: Kabol, Velayat-e

FMNH

102024

Afghanistan: Kabol, Velayat-e

FMNH

102025

Afghanistan: Kabol, Velayat-e

FMNH

102028

Afghanistan: Kabol, Velayat-e

BMNH

81.8.16.2

?

Afghanistan: Qandahar, Velayat-e

BMNH

79.11.21.515

?

Afghanistan: Qandahar, Velayat-e

BMNH

79.11.21.516

?

Afghanistan: Qandahar, Velayat-e

FMNH

102030

Afghanistan: Qandahar, Velayat-e

FMNH

102036

Afghanistan: Qandahar, Velayat-e

FMNH

102037

Afghanistan: Qandahar, Velayat-e

FMNH

102031

Afghanistan: Qandahar, Velayat-e

FMNH

102032

Afghanistan: Qandahar, Velayat-e

FMNH

102033

Afghanistan: Qandahar, Velayat-e

FMNH

102034

Afghanistan: Qandahar, Velayat-e

FMNH

102035

Afghanistan: Qandahar, Velayat-e

BMNH

69.8.24.2

Afghanistan: “Beloschittan”

USNM

240761

China: Gansu Sheng

USNM

240758

China: Gansu Sheng

USNM

240759

China: Gansu Sheng

USNM

240760

China: Gansu Sheng

BEIJ

24796

China: Nei Monggol Zizhiqu Dixing

BEIJ

18327

China: Nei Monggol Zizhiqu Dixing

BEIJ

18332

China: Nei Monggol Zizhiqu Dixing

BEIJ

21918

China: Nei Monggol Zizhiqu Dixing

BMNH

75.3.30.5

China: Xinjiang Uygur Zizhiqu Dixing

USNM

62181

China: Xinjiang Uygur Zizhiqu Dixing

USNM

62184

China: Xinjiang Uygur Zizhiqu Dixing

BEIJ

24795

China: Xinjiang Uygur Zizhiqu Dixing

BEIJ

10659

China: Xinjiang Uygur Zizhiqu Dixing

BEIJ

10662

China: Xinjiang Uygur Zizhiqu Dixing

BEIJ

13610

China: Xinjiang Uygur Zizhiqu Dixing

BEIJ

13611

China: Xinjiang Uygur Zizhiqu Dixing

BEIJ

13612

China: Xinjiang Uygur Zizhiqu Dixing

BEIJ

21236

China: Xinjiang Uygur Zizhiqu Dixing

BEIJ

13608

China: Xinjiang Uygur Zizhiqu Dixing

BEIJ

24794

China: Xinjiang Uygur Zizhiqu Dixing

BEIJ

23269

China: Xinjiang Uygur Zizhiqu Dixing

BEIJ

13604

China: Xinjiang Uygur Zizhiqu Dixing

BEIJ

19275

China: Xinjiang Uygur Zizhiqu Dixing

BEIJ

21237

China: Xinjiang Uygur Zizhiqu Dixing

BEIJ

21753

China: Xinjiang Uygur Zizhiqu Dixing

BEIJ

13295

China: Xinjiang Uygur Zizhiqu Dixing

BEIJ

13603

China: Xinjiang Uygur Zizhiqu Dixing

BEIJ

13607

China: Xinjiang Uygur Zizhiqu Dixing

BEIJ

21238

China: Xinjiang Uygur Zizhiqu Dixing

BMNH

89.6.8.1

China: Xinjiang Uygur Zizhiqu Dixing

USNM

62182

China: Xinjiang Uygur Zizhiqu Dixing

USNM

62183

China: Xinjiang Uygur Zizhiqu Dixing

USNM

62185

China: Xinjiang Uygur Zizhiqu Dixing

BEIJ

25722

?

China: Xinjiang Uygur Zizhiqu Dixing

BEIJ

25721

?

China: Xinjiang Uygur Zizhiqu Dixing

AMNH

31246

?

China: Xinjiang Uygur Zizhiqu Dixing

BMNH

66.675

?

China: Xinjiang Uygur Zizhiqu Dixing

BMNH

75.3.30.4

?

China: Xinjiang Uygur Zizhiqu Dixing

USNM

311692

Egypt: Jizah, Mudiriyat al

USNM

311730

Egypt: Kafr ash Shaykh, Mudiriyat

USNM

311706

Egypt: Sahra al Gharbiyah, Muhafazat

USNM

311710

Egypt: Sahra al Gharbiyah, Muhafazat

USNM

311729

Egypt Sharqiyah, Mudiriyat ash

FMNH

96350

Iran: Kerman, Ostan-e

FMNH

96353

Iran: Kerman, Ostan-e

FMNH

96354

Iran: Kerman, Ostan-e

FMNH

96357

Iran: Kerman, Ostan-e

FMNH

96347

Iran: Kerman, Ostan-e

FMNH

96349

Iran: Kerman, Ostan-e

FMNH

96352

Iran: Kerman, Ostan-e

FMNH

96355

t

Iran: Kerman, Ostan-e

FMNH

96356

Iran: Kerman, Ostan-e

FMNH

96358

Iran: Kerman, Ostan-e

FMNH

96359

Iran: Kerman, Ostan-e

FMNH

96360

Iran: Kerman, Ostan-e

FMNH

96361

Iran: Kerman, Ostan-e

FMNH

96362

Iran: Kerman, Ostan-e

FMNH

96363

Iran: Kerman, Ostan-e

FMNH

96346

Iran: Khorasan, Ostan-e

USNM

326683

Iran: Khorasan, Ostan-e

USNM

326684

Iran: Khorasan, Ostan-e

USNM

326685

Iran: Khorasan, Ostan-e

USNM

326686

Iran: Khorasan, Ostan-e

USNM

326687

Iran: Khorasan, Ostan-e

USNM

326689

Iran: Khorasan, Ostan-e

USNM

326690

Iran: Khorasan, Ostan-e

USNM

326691

Iran: Khorasan, Ostan-e

USNM

326693

Iran: Khorasan, Ostan-e

USNM

326694

Iran: Khorasan, Ostan-e

USNM

326703

Iran: Khorasan, Ostan-e

USNM

326705

Iran: Khorasan, Ostan-e

USNM

327911

Iran: Khorasan, Ostan-e

FMNH

96348

Iran: Khorasan, Ostan-e

USNM

326675

Iran: Khorasan, Ostan-e

USNM

326676

Iran: Khorasan, Ostan-e

USNM

326677

Iran: Khorasan, Ostan-e

USNM

326678

Iran: Khorasan, Ostan-e

USNM

326679

Iran: Khorasan, Ostan-e

USNM

326680

Iran: Khorasan, Ostan-e

USNM

326681

Iran: Khorasan, Ostan-e

USNM

326682

Iran: Khorasan, Ostan-e

USNM

326688

Iran: Khorasan, Ostan-e

USNM

326692

Iran: Khorasan, Ostan-e

USNM

327912

Iran: Khorasan, Ostan-e

USNM

350115

Iran: Khuzestan

USNM

340934

Iran: Mazanderan, Ostan-e

USNM

340936

Iran: Mazanderan, Ostan-e

USNM

340938

Iran: Mazanderan, Ostan-e

USNM

340933

Iran: Mazanderan, Ostan-e

USNM

340933

Iran: Mazanderan, Ostan-e

AMNH

184605

?

Israel

AMNH

57221

Mongolia: Hovd Aymag

AMNH

57216

Mongolia: Hovd Aymag

AMNH

57217

Mongolia: Hovd Aymag

AMNH

57222

Mongolia: Hovd Aymag

HMCZ

20683

Mongolia: Hovd Aymag

AMNH

34001

Mongolia: Ovorhangay Aymag

AMNH

217232

?

Pakistan: Baluchistan Province

AMNH

217233

?

Pakistan: Baluchistan Province

AMNH

217234

?

Pakistan: Baluchistan Province

AMNH

217235

?

Pakistan: Baluchistan Province

AMNH

244377

?

Pakistan: Baluchistan Province

AMNH

244378

?

Pakistan: Baluchistan Province

AMNH

244379

?

Pakistan: Baluchistan Province

AMNH

244380

?

Pakistan: Baluchistan Province

AMNH

244381

?

Pakistan: Baluchistan Province

AMNH

244382

?

Pakistan: Baluchistan Province

AMNH

244383

?

Pakistan: Baluchistan Province

BMNH

82.883

?

Pakistan: Baluchistan Province

BMNH

82.884

?

Pakistan: Baluchistan Province

AMNH

170230

Pakistan: Baluchistan Province

BMNH

19.11.8.6

Pakistan: Baluchistan Province

BMNH

19.11.7.18

Pakistan: Baluchistan Province

FMNH

82613

Pakistan: Baluchistan Province

USNM

368929

Pakistan: Baluchistan Province

AMNH

170226

Pakistan: Baluchistan Province

AMNH

170227

Pakistan: Baluchistan Province

AMNH

170228

Pakistan: Baluchistan Province

AMNH

170229

Pakistan: Baluchistan Province

AMNH

244384

Pakistan: Baluchistan Province

BMNH

19.11.8.4

Pakistan: Baluchistan Province

BMNH

19.11.8.5

Pakistan: Baluchistan Province

FMNH

82612

Pakistan: Baluchistan Province

USNM

200318

Pakistan: Baluchistan Province

USNM

410930

Pakistan: Baluchistan Province

USNM

413464

Pakistan: Baluchistan Province

AMNH

31248

?

Palestine: Sarepta (= as Sarafand, Lebanon?)

BMNH

7.1.1.8

?

U.S.S.R.: “Volga”

BMNH

7.1.1.129

?

U.S.S.R.: “Volga”

AMNH

85309

U.S.S.R.: Kazakh.S.S.R., KzylOrdinsk.

AMNH

85308

U.S.S.R.: Kazakh.S.S.R., KzylOrdinsk.

USNM

258739

?

U.S.S.R.: Kazakh.S.S.R., Akmolinsk.

BMNH

11.12.14.5

U.S.S.R.: Kazakh.S.S.R., TaldyKurgansk.

BMNH

14.5.10.31

U.S.S.R.: Kazakh.S.S.R., TaldyKurgansk.

BMNH

14.5.10.32

U.S.S.R.: Kazakh.S.S.R., TaldyKurgansk.

BMNH

11.12.14.3

U.S.S.R.: Kazakh.S.S.R., TaldyKurgansk.

BMNH

11.12.14.1

U.S.S.R.: Kazakh.S.S.R., TaldyKurgansk.

BMNH

11.12.14.2

U.S.S.R.: Kazakh.S.S.R., TaldyKurgansk.

BMNH

11.12.14.4

U.S.S.R.: Kazakh.S.S.R., TaldyKurgansk.

USNM

251642

?

U.S.S.R.: R.S.F.S.R., Altaisk. krai

HMCZ

23298

?

U.S.S.R.: R.S.F.S.R., Altaisk. krai

HMCZ

23299

?

U.S.S.R.: R.S.F.S.R., Altaisk. krai

BMNH

20.7.4.1

?

U.S.S.R.: Turkmen.S.S.R., Krasnovodsk.

FMNH

106648

U.S.S.R.: Turkmen.S.S.R., Krasnovodsk.

KU

117036

?

U.S.S.R.: Turkmen.S.S.R., Maryisk.

AMNH

176282

U.S.S.R.: Turkmen.S.S.R., Maryisk.

AMNH

87085

?

U.S.S.R.: Uzbek.S.S.R., Fergansk.

Hemiechinus collaris

HMCZ

5187

?

India

AMNH

214190

India

FMNH

−

?

Pakistan: Punjab Province

FMNH

−

Pakistan: Punjab Province

USNM

326277

?

Pakistan: Punjab Province

USNM

326278

Pakistan: Punjab Province

AMNH

185103

?

Pakistan: Sind Province

BMNH

75.06

Pakistan: Sind Province

BMNH

75.09

Pakistan: Sind Province

BMNH

75.1399

Pakistan: Sind Province

BMNH

75.1400

Pakistan: Sind Province

BMNH

75.05

Pakistan: Sind Province

BMNH

75.08

Pakistan: Sind Province

BMNH

80.12.15.11

Pakistan: Sind Province

Hemiechinus hypomelas

FMNH

102038

Afghanistan: Nangarhar, Velayat-e

FMNH

102039

Afghanistan: Nangarhar, Velayat-e

BMNH

81.8.16.3

?

Afghanistan: Qandahar, Velayat-e

BMNH

85.11.5.5

Afghanistan: Qandahar, Velayat-e

USNM

327914

Iran: Dasht-e-lut

USNM

327915

Iran: Dasht-e-lut

USNM

327913

Iran: Dasht-e-lut

FMNH

110959

?

Iran: Fars, Ostan-e

FMNH

57974

?

Iran: Fars, Ostan-e

FMNH

96365

?

Iran: Fars, Ostan-e

FMNH

110960

Iran: Fars, Ostan-e

FMNH

110961

?

Iran: Fars, Ostan-e

FMNH

110962

?

Iran: Fars, Ostan-e

FMNH

110063

?

Iran: Flam, Ostan-e

USNM

326701

Iran: Khorasan, Ostan-e

USNM

326697

Iran: Khorasan, Ostan-e

USNM

326695

Iran: Khorasan, Ostan-e

USNM

326699

Iran: Khorasan, Ostan-e

USNM

326696

Iran: Khorasan, Ostan-e

USNM

326698

Iran: Khorasan, Ostan-e

USNM

326700

Iran: Khorasan, Ostan-e

BMNH

21.12.3.2

Iran: Tanb Island

BMNH

88.6.18.3

?

Oman: Muscat

BMNH

85.11.5.5

Oman: Muscat

USNM

352951

Pakistan: Baluchistan Province

USNM

368930

Pakistan: Baluchistan Province

FMNH

82609

?

Pakistan: Punjab Province

FMNH

82608

Pakistan: Punjab Province

USNM

368933

Pakistan: Punjab Province

USNM

368937

Pakistan: Punjab Province

USNM

368931

Pakistan: Punjab Province

USNM

368932

Pakistan: Punjab Province

USNM

368934

Pakistan: Punjab Province

USNM

368935

Pakistan: Punjab Province

USNM

368936

Pakistan: Punjab Province

USNM

410929

Pakistan: Punjab Province

BMNH

87.4.2.2

Pakistan: Sind Province

BMNH

2.11.22.2

Yemen (Aden)

Hemiechinus micropus

FMNH

82604

India

FMNH

82605

India

FMNH

82607

India

FMNH

82606

India

USNM

369316

Pakistan: Punjab Province

KU

125480

?

Pakistan: Sind Province

BMNH

75.1413

Pakistan: Sind Province

BMNH

75.1418

Pakistan: Sind Province

BMNH

75.1414

Pakistan: Sind Province

BMNH

75.1415

Pakistan: Sind Province

BMNH

75.1416

Pakistan: Sind Province

BMNH

75.1417

Pakistan: Sind Province

Hylomys hainanensis

BEIJ

70875

China: Hainan Dao

BEIJ

70891

China: Hainan Dao

BEIJ

70892

China: Hainan Dao

bEIj

70949

China: Hainan Dao

BEIJ

70886

China: Hainan Dao

BEIJ

70904

China: Hainan Dao

BEIJ

70929

China: Hainan Dao

Hylomys sinensis

AMNH

115505

Burma: Kachin State

AMNH

115507

Burma: Kachin State

AMNH

115508

Burma: Kachin State

AMNH

115511

Burma: Kachin State

AMNH

115513

Burma: Kachin State

AMNH

115514

Burma: Kachin State

AMNH

115515

Burma: Kachin State

AMNH

115516

Burma: Kachin State

AMNH

115522

Burma: Kachin State

AMNH

115503

Burma: Kachin State

AMNH

115504

Burma: Kachin State

AMNH

115506

Burma: Kachin State

AMNH

115509

Burma: Kachin State

AMNH

115510

Burma: Kachin State

AMNH

115512

Burma: Kachin State

AMNH

115517

Burma: Kachin State

AMNH

115518

Burma: Kachin State

AMNH

115519

Burma: Kachin State

AMNH

115520

Burma: Kachin State

AMNH

115521

Burma: Kachin State

AMNH

115523

Burma: Kachin State

AMNH

115524

Burma: Kachin State

AMNH

115525

Burma: Kachin State

BMNH

9.12.13.1

?

China: Sichuan Sheng

USNM

258124

?

China: Sichuan Sheng

USNM

258125

?

China: Sichuan Sheng

USNM

258126

?

China: Sichuan Sheng

USNM

258127

?

China: Sichuan Sheng

USNM

258129

?

China: Sichuan Sheng

BMNH

17.2.1.19

China: Sichuan Sheng

BMNH

17.2.1.20

China: Sichuan Sheng

FMNH

36170

China: Sichuan Sheng

FMNH

37017

China: Sichuan Sheng

FMNH

37020

China: Sichuan Sheng

FMNH

37021

China: Sichuan Sheng

FMNH

37024

China: Sichuan Sheng

FMNH

39664

China: Sichuan Sheng

FMNH

39665

China: Sichuan Sheng

FMNH

39666

China: Sichuan Sheng

FMNH

39667

China: Sichuan Sheng

USNM

258128

China: Sichuan Sheng

BMNH

17.2.1.14

;

China: Sichuan Sheng

BMNH

17.2.1.15

China: Sichuan Sheng

BMNH

17.2.1.16

China: Sichuan Sheng

BMNH

17.2.1.17

China: Sichuan Sheng

BMNH

17.2.1.18

China: Sichuan Sheng

FMNH

37016

China: Sichuan Sheng

FMNH

37018

China: Sichuan Sheng

FMNH

37019

China: Sichuan Sheng

FMNH

37022

China: Sichuan Sheng

FMNH

37023

China: Sichuan Sheng

BMNH

11.8.6.1

?

China: Yunnan Sheng

AMNH

44235

China: Yunnan Sheng

AMNH

44239

China: Yunnan Sheng

AMNH

44240

China: Yunnan Sheng

AMNH

44242

China: Yunnan Sheng

AMNH

44245

China: Yunnan Sheng

AMNH

44248

China: Yunnan Sheng

AMNH

44254

China: Yunnan Sheng

AMNH

44261

China: Yunnan Sheng

AMNH

44262

China: Yunnan Sheng

AMNH

44263

China: Yunnan Sheng

AMNH

44270

China: Yunnan Sheng

BMNH

12.7.15.1

China: Yunnan Sheng

FMNH

35776

China: Yunnan Sheng

FMNH

35777

China: Yunnan Sheng

FMNH

35778

China: Yunnan Sheng

FMNH

35779

China: Yunnan Sheng

FMNH

35780

China: Yunnan Sheng

FMNH

35782

China: Yunnan Sheng

FMNH

35783

China: Yunnan Sheng

HMCZ

20692

China: Yunnan Sheng

USNM

241428

China: Yunnan Sheng

AMNH

44244

China: Yunnan Sheng

AMNH

44249

China: Yunnan Sheng

AMNH

44255

China: Yunnan Sheng

AMNH

44256

China: Yunnan Sheng

AMNH

44260

China: Yunnan Sheng

AMNH

44264

China: Yunnan Sheng

AMNH

44265

China: Yunnan Sheng

AMNH

44267

China: Yunnan Sheng

AMNH

44268

China: Yunnan Sheng

AMNH

44271

China: Yunnan Sheng

BMNH

14.10.23.3

China: Yunnan Sheng

FMNH

35775

China: Yunnan Sheng

FMNH

35781

China: Yunnan Sheng

FMNH

35784

China: Yunnan Sheng

HMCZ

20688

China: Yunnan Sheng

HMCZ

20690

China: Yunnan Sheng

HMCZ

20691

China: Yunnan Sheng

HMCZ

20689

China: Yunnan Sheng

USNM

241402

China: Yunnan Sheng

USNM

258338

?

Malaysia: Sabah

FMNH

38889

?

Vietnam: Hoang Lien Son, Tinh

FMNH

41297

?

Vietnam: Hoang Lien Son, Tinh

FMNH

41298

?

Vietnam: Hoang Lien Son, Tinh

FMNH

41299

?

Vietnam: Hoang Lien Son, Tinh

FMNH

38887

Vietnam: Hoang Lien Son, Tinh

FMNH

38888

Vietnam: Hoang Lien Son, Tinh

HMCZ

27047

Vietnam: Hoang Lien Son, Tinh

HMCZ

38262

Vietnam: Hoang Lien Son, Tinh

FMNH

38890

Vietnam: Hoang Lien Son, Tinh

FMNH

38891

Vietnam: Hoang Lien Son, Tinh

FMNH

38892

Vietnam: Hoang Lien Son, Tinh

FMNH

38893

Vietnam: Hoang Lien Son, Tinh

FMNH

38894

Vietnam: Hoang Lien Son, Tinh

FMNH

38895

Vietnam: Hoang Lien Son, Tinh

FMNH

38896

Vietnam: Hoang Lien Son, Tinh

FMNH

38897

Vietnam: Hoang Lien Son, Tinh

FMNH

38898

Vietnam: Hoang Lien Son, Tinh

FMNH

41296

Vietnam: Hoang Lien Son, Tinh

HMCZ

38252

Vietnam: Hoang Lien Son, Tinh

HMCZ

38261

Vietnam: Hoang Lien Son, Tinh

Hylomys suillus

USNM

481285

Indonesia: Java

HMCZ

20687

Burma: Kachin State

AMNH

44113

China: Yunnan Sheng

AMNH

44272

China: Yunnan Sheng

AMNH

44112

China: Yunnan Sheng

AMNH

44275

China: Yunnan Sheng

FMNH

32309

China: Yunnan Sheng

USNM

481289

Indonesia: Java

USNM

521660

Indonesia: Java

USNM

521661

Indonesia: Java

USNM

481286

Indonesia: Java

AMNH

87313

Laos: Bolovens, Plateau des

FMNH

32308

Laos: Phong Saly, Khoueng

BMNH

26.10.4.42

Laos: Xien Quang Koo

USNM

292338

?

Malaysia: Sabah

USNM

292339

?

Malaysia: Sabah

USNM

292340

?

Malaysia: Sabah

USNM

292356

Malaysia: Sabah

USNM

292357

Malaysia: Sabah

USNM

292348

Malaysia: Sabah

USNM

292354

Malaysia: Sabah

USNM

300816

Malaysia: Sabah

USNM

487911

Malaysia: West

USNM

307548

Malaysia: West

USNM

487920

Malaysia: West

BMNH

20.7.3.12

Thailand

HMCZ

35452

Thailand: Chiang Mai Changwat

HMCZ

35453

Thailand: Chiang Mai Changwat

USNM

320486

Vietnam

USNM

320492

Vietnam

BMNH

26.10.4.40

Vietnam: Gia Lai-Kon Turn, Tinh

BMNH

26.10.4.41

Vietnam: Gia Lai-Kon Turn, Tinh

BMNH

26.10.4.36

Vietnam: Gia Lai-Kon Turn, Tinh

BMNH

26.10.4.37

Vietnam: Gia Lai-Kon Turn, Tinh

BMNH

26.10.4.38

Vietnam: Gia Lai-Kon Turn, Tinh

BMNH

26.10.4.39

Vietnam: Gia Lai-Kon Turn, Tinh

FMNH

46631

Vietnam: Lam Dong, Tinh

FMNH

46632

Vietnam: Lam Dong, Tinh

FMNH

46633

Vietnam: Lam Dong, Tinh

FMNH

46634

Vietnam: Lam Dong, Tinh

FMNH

46636

d

Vietnam: Lam Dong, Tinh

FMNH

46637

d

Vietnam: Lam Dong, Tinh

FMNH

46638

d

Vietnam: Lam Dong, Tinh

Mesechinus dauuricus

BEIJ

01826

China: Jilin Sheng Dixing

BEIJ

01827

China: Jilin Sheng Dixing

BEIJ

01828

China: Jilin Sheng Dixing

MGU

83740

China: Liaoning Sheng

USNM

270540

China: Liaoning Sheng

USNM

270539

China: Liaoning Sheng

BEIJ

21295

China: Nei Monggol Zizhiqu Dixing

BEIJ

24276

China: Nei Monggol Zizhiqu Dixing

BEIJ

10666

China: Nei Monggol Zizhiqu Dixing

BEIJ

15855

China: Nei Monggol Zizhiqu Dixing

BEIJ

20565

China: Nei Monggol Zizhiqu Dixing

BEIJ

24277

China: Nei Monggol Zizhiqu Dixing

BEIJ

25210

China: Nei Monggol Zizhiqu Dixing

BEIJ

10667

China: Nei Monggol Zizhiqu Dixing

BEIJ

10665

China: Nei Monggol Zizhiqu Dixing

BEIJ

20842

China: Nei Monggol Zizhiqu Dixing

BMNH

9.1.1.2

China: Shaanxi Sheng

BMNH

9.1.1.3

China: Shaanxi Sheng

BMNH

9.1.1.4

China: Shaanxi Sheng

BMNH

9.1.1.5

China: Shaanxi Sheng

BMNH

9.1.1.6

China: Shaanxi Sheng

BMNH

9.1.1.7

China: Shaanxi Sheng

BMNH

9.1.1.8

China: Shaanxi Sheng

BMNH

9.1.1.9

China: Shaanxi Sheng

BMNH

9.1.1.10

China: Shaanxi Sheng

MGU

107219

Mongolia: Ch'er-Khangai Aymag

MGU

130413

?

Mongolia: Khentei Aymag

MGU

113799

?

Mongolia: Vostochnyi Aymag

MGU

100455

U.S.S.R.: Buryat-A.S.S.R.

MGU

42341

U.S.S.R.: Buryat-A.S.S.R.

MGU

100456

U.S.S.R.: Buryat-A.S.S.R.

MGU

42340

?

U.S.S.R.: R.S.F.S.R. Chitinsk.

MGU

83733

U.S.S.R.: R.S.F.S.R. Chitinsk.

MGU

83738

U.S.S.R.: R.S.F.S.R. Chitinsk.

MGU

43833

U.S.S.R.: R.S.F.S.R. Chitinsk.

MGU

83732

U.S.S.R.: R.S.F.S.R. Chitinsk.

MGU

83735

U.S.S.R.: R.S.F.S.R. Chitinsk.

MGU

83737

U.S.S.R.: R.S.F.S.R. Chitinsk.

Mesechinus hughi

BMNH

0.6.27.2

China: Shaanxi Sheng

BEIJ

19434

China: Shaanxi Sheng

MGU

83744

China: Shaanxi Sheng

MGU

83745

China: Shaanxi Sheng

BEIJ

19433

China: Shaanxi Sheng

Microgale talaiaci

USNM

341696

Madagascar

Podogymnura aureospinula

UPLB

3753

Philippine Islands: Dinagat Island

Podogymnura truei

DMNH

5948

?

Philippine Islands: Mindanao Island

DMNH

5949

Philippine Islands: Mindanao Island

DMNH

5950

Philippine Islands: Mindanao Island

DMNH

5951

Philippine Islands: Mindanao Island

DMNH

5952

Philippine Islands: Mindanao Island

DMNH

5953

Philippine Islands: Mindanao Island

KU

98141

Philippine Islands: Mindanao Island

Setifer setosus

USNM

122489

?

Madagascar

USNM

254983

?

Madagascar

Solenodon cubanus

USNM

2230/1395

?

Cuba

Solenodon paradoxus

USNM

290520

Zoo Specimen

USNM

364952

Zoo Specimen

Tenrec ecaudatus

USNM

294495

?

Madagascar

USNM

63313

Madagascar

Uropsilus andersoni

ANSP

16594

China: Sichuan Sheng

Uropsilus soricipes

USNM

256119

?

China: Sichuan Sheng

USNM

260743

?

China: Sichuan Sheng

USNM

260751

China: Sichuan Sheng

Appendix 2

Data Matrix

Abbreviations arc as follows:

ANCE = Hypothetical ancestor

HEMH = Hemiechinus hypomelas

ATXF = Atelerixfrontalis

ECHG = Echinosorex gymnura

HEMM = Hemiechinus micropus

ATXG = Atelerix algirus

PODA = Podogymnura aureospinula

HEMA = Hemiechinus auritus

ATXA = Atelerix albiventris

PODT = Podogymnura truei

HEMC = Hemiechinus collaris

ATXS = Atelerix sclateri

HYLS = Hylomys sinensis

MESD = Mesechinus dauuricus

TENR = Tenrecoids

HYLU = Hylomys suillus

ERIA = Erinaceus amurensis

SORI = Soricoids

HYLH = Hylomys hainanensis

ERIC = Erinaceus concolor

LEPT = †Leptictidae

HEME = Hemiechinus aethiopicus

ERIE = Erinaceus europaeus

Unordered characters are 10, 20, 49, 62, 77, 79.

Unpolarized characters are 14, 54, 58, 61, 70, 71, 72.

Appendix 3

Apomorphy List by Stems

Abbreviations are explained in Appendix 2. Double daggers (‡) denote characters of analytically ambiguous placement.

Unordered characters are 10, 20, 49, 62, 77, 79.

Unpolarized characters are 14, 54, 58, 61, 70, 71, 72.

Appendix 4

Changes on Stems by Transformation Series

Unordered or unpolarized transformations are noted with a “U.” Characters of ambiguous placement are noted by a double dagger (‡).

Literature Cited

Agrawal, V.C.

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Plates 1–19

PLATE 1.—Echinosorex gymnura, USNM 487891, . Malaysia: Selangor, Klang; Tanjong Duablas, Kuala Langat Forest Reserve, Bukit Mandol (2°55′N,101°34′E). CBL = 73.45 mm.

PLATE 2.—Podogymnura aureospinula, UPBL 3753, . Philippine Islands: Dinagat Island; Loreto Municipality, Balitbiton. CBL = 52.02 mm.

PLATE 3.—Podogymnura truei, KU 98141, . Philippine Islands: Mindinao Island. CBI. = 38.25 mm.

PLATE 4.—Hylomys hainanensis, BELT 70875, . China: Hainan Dao. CBL = 34.82 mm.

PLATE 5.—Hylomys sinensis, USNM 241402, . China: Yunnan; Ho Mu Shu Pass, on main Yung-Chang to Teng-yueh Road. CBL = 28.34 mm.

PLATE 6.—Hylomys suillus, USNM 292347, . Malaysia: Sabah; Mount Kinabalu; Bundu Tuhan. CBL = 34.00 mm.

PLATE 7.—Atelerix albiventris, USNM 181442, . Kenya: South Nguasso Nyiro; Loita Plains. CBL = 40.20 mm.

PLATE 8.—Atelerix algirus, USNM 476058, . Morocco: Agadir, 8 km N of Tamri (30°20′N, 9°50′W). CBL = 57.62 mm.

PLATE 9.—Atelerix frontalis, USNM 267653, . South Africa: Transvaal; Pretoria District. CBL = 47.41 mm.

PLATE 10.—Atelerix sclateri, USNM 63220, . Somaliland: Berbera. CBL = 39.85 mm.

PLATE 11.—Erinaceus amurensis, USNM 197779, ;. China: Kirin [Manchuria]; 60 mi SW on Sungaree River. CBL= 48.80 mm.

PLATE 12.—Erinaceus concolor. USNM 369533, . Iran: Khuzistan; Dorad, 5 km NW. CBL = 58.91 mm.

PLATE 13.—Erinaceus europaeus, USNM I53410, . United Kingdom: England; Surrey. CBL = 55.70 mm.

PLATE 14.—Hemiechinus aethiopicus, USNM 470566, Morocco: Ouarzazate; Tazarine, 3 km NW (30°48′N, 5°36′W). CBL = 48.90 mm.

PLATE 15.—Hemiechinus auritus, USNM 340933, . Iran: Mazandcran; 28 km W of Gorgan, on Su River near Ghararch. CBL = 49.40 mm.

PLATE 16.—Hemiechinus collaris, FMNH 82611, . India: Gujerat. CBL = 47.40 mm.

PLATE 17.—Hemiechinus hypomelas, USNM 326697, . Iran: Khurasan; Turbat-i-Haidari. CBL = 54.90 mm.

PLATE 18.—Hemiechinus micropus, FMNH 82605, . India: Kathiawar. CBL = 44.60 mm.

PLATE 19.—Mesechinus dauuricus, USNM 270539, . China: Inner Mongolia; Northern Manchuria; Barga, Dalai Nor Lake at the mouth of the Urshun River. CBL = 49.65 mm.

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Text-reference system (author, year:page used within the text, with full citation in “Literature Cited” at the end of the text) must be used in place of bibliographic footnotes in all Contributions Series and is strongly recommended in the Studies Series: “(Jones. 1910:122)” or “…Jones (1910:122)” If bibliographic footnotes are required, use the short form (author, brief title, page) with the full citation in the bibliography.

Footnotes, when few in number, whether annotative or bibliographic, should be typed on separate sheets and inserted immediately after the text pages on which the references occur. Extensive notes must be gathered together and placed at the end of the text in a notes section.

Bibliography, depending upon use, is termed “Literature Cited,” “References,” or “Bibliography.” Spell out titles of books, articles, journals, and monographic series. For book and article titles use sentence-style capitalization according to the rules of the language employed (exception: capitalize all major words in English), For journal and series titles, capitalize the initial word and all subsequent words except articles, conjunctions, and prepositions. Transliterate languages that use a non-Roman alphabet according to the Library of Congress system. Underline (for italics) titles of journals and series and titles of books that are not part of a series. Use the parentheses/colon system for volume (number): pagination: “10(2):5–9.” For alignment and arrangement of elements, follow the format of recent publications in the series for which the manuscript is intended. Guidelines for preparing bibliography may be secured from Series Section, SI Press

Legends for illustrations must be submitted at the end of the manuscript, with as many legends typed, double-spaced, to a page as convenient.

Illustrations must be submitted as original art (not copies) accompanying, but separate from, the manuscript. Guidelines for preparing art may be secured from Series Section. SI Press All types of illustrations (photographs, line drawings, maps, etc.) may be intermixed throughout the printed text They should be termed Figures and should be numbered consecutively as they will appear in the monograph. If several illustrations are treated as components of a single composite figure, they should be designated by lowercase italic letters on the illustration; also, in the legend and in text references the italic letters (underlined in copy) should be used: “Figure 9b.” Illustrations that are intended to follow the printed text may be termed Plates, and any components should be similarly lettered and referenced: “Plate 9b” Keys to any symbols within an illustration should appear on the art rather than in the legend.

Some points of style: Do not use periods after such abbreviations as “mm, ft, USNM, NNE.” Spell out numbers “one” through “nine” in expository text, but use digits in all other cases if possible. Use of the metric system of measurement is preferable; where use of the English system is unavoidable, supply metric equivalents in parentheses. Use the decimal system for precise measurements and relationships, common fractions for approximations. Use day/month/year sequence for dates: “9 April 1976.” For months in tabular listings or data sections, use three-letter abbreviations with no periods: “Jan. Mar, Jun.” etc. Omit space between initials of a personal name: “J B. Jones.”

Arrange and paginate sequentially every sheet of manuscript in the following order: (1) title page. (2) abstract, (3) contents, (4) foreword and/or preface, (5) text, (6) appendixes, (7) notes section, (8) glossary, (9) bibliography. (10) legends, (11) tables. Index copy may be submitted at page proof stage, but plans for an index should be indicated when manuscript is submitted.