AMERICAN MUSEUM Novtitates
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Number 3340, 52 pp., 9 figures, 2 tables, 6 appendices June 22, 2001
The Phylogenetic Resolving Power of Discrete Dental Morphology Among Extant Hedgehogs and the Implications for Their Fossil Record
GINA C. GOULD!
CONTENTS
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DD Analysis 2: Covariation of Polymorphism and Sexual Dimorphism .......... 14
DD Analysis 3: Covariation of Polymorphism and Relative Age ................ 14
DD Analysis 4: Covariation of Polymorphism and Geographic Locality .......... 14 FADE 14 05 VIO SIS are wcysestaeiieot ts tate Hee ek eee D Ce wi segh ee sleiemaeel ee Bah hus ve Ot eee Rate tact ey 16 PSO SEM ete A MALY RRS co cele ence Yordee t we eirtoskeced UV ie hind he ste eile eee te ed eatierd ob FT ae nag 16 Methods aiid SUS tC Attn oa ecg eet ees YEA Pat Saxton 5. «murs Dad eee, PIII IP Pigg eon. + + 16
‘Graduate Student, Division of Paleontology, American Museum of Natural History; present address: Florida Museum of Natural History, University of Florida, RO. Box 112710, Gainesville, FL 32641.
Copyright © American Museum of Natural History 2001 ISSN 0003-0082 / Price $5.80
pe AMERICAN MUSEUM NOVITATES NO. 3340
IRCSUNS ie 3 ae Cet Re ee Bh PEL ASE bathe Lew ee EI det! sears $1 BOO 17 Phylogenetic: Anadlysisslata: Sets 1 ey ee ae Te BT ee tok ere ee ala 17 Phylogenctic“Analysis2r Data, Sete2-a rs Atveeih sia hes pa ven aos neste ae Se pha we 17 BricleS WnOpSis— 4.45737 oie oss Se Baler th es AP ver eee CE a ea ew Lecelokns s oo wien EE 18
DPS OUSSIGT Ae EN or ool ee lk Meo de ewe ks tay) MOR SPAE ND Sens 2 at ee tee ep ney, SoRW Meats cameo son Se 18 Corchiston 5A iA Oe UC Pde 8 UR al A Fee Se REY 3 Stl ae, 21 reN(od aca nud (exe 24 LOU cd 01 kc Ew egeONOay We PaPO JN PNGR Be cope Ep ane ay See CnC AP Re PaO EN ON Sy oo ae satan CE nS Pr PP oA PRETSLOTIC ES i eB acta eel gs sled ad Somers ee ek Bes eangs he et) AS Ask ag eee, Rts os, Gi ke teE faz APPENDIX 12 ‘Transformation. Series. Considered: 42.0.0. on ae a ee See 28 APPENDIXAAZ SSpeCnnens IReVIC WER | 4 5.5 b Rec ata 53 oeakes & & OE aa 3 3 ape ER Rath isis Rags & OE 33 APPENDIX 3 Brequéney :DistributiOi: gusc< a we tea bt ee ee Pees eek EEE Ss 36 APPENDIX 4: Transformation Series Recovered for Phylogenetic Review ............ 45 APPENDIX. 5: Phylogenetic Analysis of Data,Set. A cc46 3. ee eee ee bh 48
APPENDIX 6+Phylogénetic Analysis: of Data-Set Ber: fers vs oe Meeting $4 eee BEES 50
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GOULD: DENTAL MORPHOLOGY OF HEDGEHOGS
ABSTRACT
Discrete dental morphology among members of the extant Erinaceidae (Mammalia; Lipo- typhla) is comprehensively reviewed in order to ascertain its phylogenetic resolving power. This analysis responds to the need to better understand the nature of the characters—discrete dental morphology—most commonly used to diagnose erinaceid fossil taxa, and reconstruct their evolutionary histories. This investigation attempts to set the parameters for a phylogenetic analysis of both fossil and living erinaceids.
The first phase of this investigation reviews 246 descriptive discrete dental transformation series—the majority of which were gathered primarily from the literature and are (or have been) considered apomorphies at various taxonomic levels within the family Erinaceidae. These characters were reviewed across 10 species of hedgehogs: a minimum of two species per extant genus (excluding the rare species), of which all are represented by series of indi- viduals. The data were compiled and analyzed for each individual for inter- and intraspecific variation (including asymmetry), and its possible covariation with sex, relative age (based on tooth eruption and wear stage), and geographic location.
The second phase tests the phylogenetic resolving power of the discrete dental transfor- mation series when considered as the sole body of evidence for hypotheses of evolutionary relationships. The discovered phylogenies of parsimony analyses of the discrete dental data are compared to previous hypotheses of relationships based on all known morphological ev- idence.
Results suggest that dental variation is intemperant both inter- and intraspecifically within the Erinaceidae and cannot unequivocally be attributed to any one of the variables considered (see above); and, more specifically, the phylogenetic resolving power of the dental data (across the considered taxa) is contingent on the inclusion of other data (1.e., cranial and postcranial material). Consequently, the applicability of this character set to the erinaceid fossil record as
the sole source of evidence for phylogenetic inference is challenged.
INTRODUCTION
This study investigates the appropriateness of using the dental morphology of hedgehogs (Mammalia; Lipotyphla; Erinaceidae) as the sole character suite for positing phylogenetic relationships in living and in fossil taxa. More specifically, it is designed to ascertain the extent of inter- and intraspecific dental variation among the living members of this group, and to determine whether interspecific variation can be correlated to the age, sex, and/or geographic locality of the reviewed individuals. The phylogenetic resolving pow- er of these data across the extant taxa will then be explored under the tenets of the par- simony principle. Subsequently, the applica- bility of these data as the sole source of ev- idence for inferring evolutionary relation- ships among the fossil taxa will be reevalu- ated.
OVERVIEW
Historically, the reliance upon discrete dental data as evidence for positing phylo- genetic relationships has been incongruous
between the extant and fossil erinaceid taxa. That is, fossil taxa are predominantly repre- sented by teeth, either isolated or in incom- plete jaws and maxillary bone. Consequent- ly, many fossil species are diagnosed and their phylogenetic histories reconstructed based almost exclusively on presumed dis- crete dental apomorphies (de Blainville, 1840; Matthew, 1903; Koerner, 1940; Htiir- zeler, 1944; Simpson, 1945; Butler, 1948, 1956a, 1956b, 1972, 1988; Crusafont et al., 1955; Friant, 1961; Van Valen, 1967; Mc- Kenna and Holton, 1967; Rich and Rich, 1971; Rich and Rasmussen, 1973; Gilbert, 1975; Krishtalka, 1976; Schwartz and Krish- talka, 1976; Krishtalka and West, 1977; Ste- vens, 1977; Black et al., 1980; Munthe and West, 1980; Engesser, 1972, 1979, 1980, ; Rich, 1981; Novacek, 1985; Novacek et al., 1985; see appendix 1). Phylogenies of the extant taxa, however, are based on compre- hensive morphological data sets that include pelage, cranial, and dental characters (Cor- bet, 1974, 1988; Frost et al., 1991; Storch and Qiu, 1991; Gould, 1995), as well as mo- lecular and morphometric ones (Ruedi et al.,
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1994; Ruedi and Fumagalli, 1996; Robbins and Setzer, 1985). Consequently, the dispa- rate treatment of the extant and fossil taxa has resulted in partitioned data sets: (1) den- tal characters that have been considered only for fossil taxa; and (2) more comprehensive characters that are applicable only to the ex- tant forms as a result of poor preservation in fossil taxa. A survey of the literature indi- cates there is very little character overlap be- tween data sets (see appendix | citations).
Polymorphism of discrete dental charac- ters has been reported in many of the extant hedgehogs (Woodward, 1896; Brockie, 1964; Van Valen, 1967; Harrison and Bates, 1985; Poduschka and Poduschka, 1986), the most extreme being complete absences of individ- ual teeth which seem to occur fairly frequent- ly both inter- and intraspecifically, as well as within individuals (Van Valen, 1967). These observations suggest that plasticity of these characters may occur more commonly, and possibly more globally (i.e., across all taxo- nomic levels), than was previously believed. A cursory review of the literature turned up 31 citations on tooth anomalies within bats, rodents, cervids, carnivores, and other lipo- typhlans (Palmer, 1937; Hall, 1940; Hooper, 1946; Kurten, 1953, 1982; Hooper, 1957; Jones, 1957; Setzer, 1957; Meester, 1959; Van Valen, 1967; Haft, 1963; Martin, 1968; Wallace, 1968; Choate, 1969; Ziegler, 1971; Fish and Whitaker, 1971; Janossy and Schmidt, 1975; Smith, 1977; Dippenaar, 1978; Woloszyn, 1978; Hall and Yalden, 1978; Nadachowski, 1978; Krausman, 1978; Beaver et al., 1982; Woods, et. al., 1982; French, 1985; Hillson, 1986; Davis, 1987; Barnosky, 1990; Jernvall, 1995; Clarke, 1997; Bell and Repenning, 1999).
Unlike many other morphological charac- ters, dental phenotype is not only a result of intrinsic (genetic and developmental) factors, it is a result of universal extrinsic factors that affect all teeth—tooth wear. Individual tooth wear patterns are a result of function (e.g., occlusal wear), diet (e.g., geographic location and/or individual preference), and sometimes idiopathic chewing behavior. Without a bet- ter understanding of the nature and frequency of dental variation, reliance on the phyloge- netic resolving power of these data can se- riously compromise any attempt to recon-
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struct a phylogeny at all levels of analysis. For example, within the Erinaceidae, the pur- ported dental apomorphies outnumber the most recent complete (nondental) morpho- logical data set (Gould, 1995) for the extant taxa by 2.5 to 1 (appendix 1), and the named fossil taxa comprise approximately 75% of the all the combined taxa at the generic level (McKenna and Bell, 1997), of which 75% are represented solely by teeth (Gould, 1995). If certain character states for a given transformation series were actually records of ontogenetic stages or discovered to be globally homoplastic (i.e., inter- and intra- specifically), hypotheses of the phylogenetic relationships of many of the fossil taxa would be rendered suspect.
Although this study focuses principally on one group of organisms and a particular data set (i.e., dental characters), the ubiquitous problem of paleontology is a paucity of ma- terial. Missing data is not an unexpected problem regardless of the taxonomic group under study or whether extant or extinct (Nixon, 1996). However, unlike the case with living taxa, the available data for many fos- sils is compromised by selective preserva- tion, and often only one type of datum is commonly preserved (e.g., vertebrae of snakes or sauropods; teeth of sharks or mam- mals; skull caps of pachycephlosaurs). As mentioned above, this phenomenon compli- cates the problem of missing data: Not only does operational missing data (i.e., missing cells in the data matrix) need to be addressed subsequent to a phylogenetic analysis (see Platnick et al., 1991; Maddison, 1993; Nixon and Carpenter, 1996), but also the ramifica- tions of the inherent missing data (i.e., the complete lack of other character sets). The effects of operational missing data can be tracked using diagnostic parsimony programs (e.g., MacClade, Clados, NONA); however, comprehensive absences of entire systems of an organism, such as skeletal or soft tissue material, pose a much more pervasive prob- lem. It seems judicious, therefore, to test the reliability of monotypic data for establishing phylogeny. That is, how much confidence can we expect to have in a phylogeny or a proposed classification that is based exclu- sively on one type of data, or simply, on one small aspect of the organism?
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Echinosorex gymnura B Podogymunura aureospinula
Podogymunura truei Aylomys sinensis Hytomys suillus Aylomys hainanensis Hemiechinus aethiopicus Hemiechinusmicropus Mioechinus Hemiechinus hypomelas Hemiechinus auritus
c Hemiechinus collaris Mesechinus dauuricus Erinaceus amurensis Erinaceus concolor Ertnaceus europaeus Alélerix frontalis Atelerix algirus Atelerix albiventris Atelerix sclateri Proterix loamisi
D Brachyerix
$>
Metechinus Neurogymmnurus Galerix
da. Lanthanotherium
Echinosorex gymnura Podogymnura aureospinula Podogymmura truei HAylomys sinensis Hylomys hainanensis Aylomys suiilus A Hemiechinus aethiopicus Hemiechinus micropus Hemiechinus hypomelas Hemiechinus auritus C Hemiechinus collaris Mesechinus dauuricus Erinaceus amurensis Frinaceus concolor Erinaceus europaeus Alelerix frontalis Alelerix algirus Alelerix albiventris b. Alelerix sclateri
Fig. 1. (a) Gould’s (1995) Adams tree, fossils are indicated in bold (b) Frost et al.’s (1991) sin- gle most parsimonious tree. A = Erinaceidae; B = Hylomyinae; C = Erinaceinae; D = Brachyer- icinae.
CURRENT TAXONOMY OF THE ERINACEIDAE
The Erinaceidae are a _ well-established monophyletic group (see fig. 1, stem A; Frost et al., 1991; Gould, 1995). There are approx- imately 19-23 reported living species, and over 30 recognized fossil genera (McKenna and Bell, 1997). A recent phylogenetic anal- ysis of both fossil and extant taxa indicates that this lineage may extend as far back as the late Cretaceous (Gould, 1995), making this group one of the oldest surviving line- ages of placental mammals.
Hypotheses of the historical relationships
GOULD: DENTAL MORPHOLOGY OF HEDGEHOGS 5
within the Erinaceidae are based almost ex- clusively on morphological data (Butler, 1948, 1988; Rich, 1981; Novacek, 1985; Frost et al., 1991; Gould, 1995; McKenna and Bell, 1997), although recently, molecular data have been employed in phylogenetic re- constructions of more inclusive groups (Rue- di et al., 1994; Ruedi and Fumagalli, 1996; Filippucci and Simson, 1996; Surin et al., 1997), and one morphometric-based phylog- eny has been proposed for the living genera (Robbins and Setzer, 1985). Thus far, how- ever, these data sets used to infer phylogeny have remained distinct, a practice that has re- sulted in incongruent hypotheses of relation- ships (see phylogenies proposed by Butler, 1948, 1988; Rich, 1981; Robbins and Setzer, 1985; Frost et al., 1991). Recent efforts to reconcile some of the disparate data sets (Ruedi et al., 1994; Gould, 1995, 1997) yielded, not surprisingly, conflicting results with all previous hypothesis of relationships that are based solely on partitioned data sets.
In the most recently proposed classifica- tion of all the known erinaceids (McKenna and Bell, 1997), four subfamilies are recog- nized (fig. 1), two of which include all extant members of the family: (1) the Hylomyinae (moonrats, or gymnures; stem B) of Malaysia and Indonesia, whose fossil record is cur- rently challenged (Gould, 1995); and (2) the Erinaceinae (spiny hedgehogs, stem C), a group distributed throughout Europe, Asia, and Africa, whose fossil members are known from all three of these regions as well as North America. The remaining two subfam- ilies, the Brachyericinae (fig. 1b, stem D) and Tupaiodontinae (not shown in fig. 1), are ex- clusively composed of fossil taxa from both North America and Asia.
DISCRETE DENTAL DATA ANALYSES METHODS AND JUSTIFICATION
CHARACTERS REVIEWED AND SOURCES OF DIFFICULTY: As mentioned in the Introduc- tion, many of the fossil taxa are represented only by isolated teeth and jaws. The majority of the characters reviewed in this analysis (see appendix 1) were gathered primarily from the paleontological literature. In addi- tion, some new characters and character states were added from personal observations
6 AMERICAN MUSEUM NOVITATES
made during the course of this analysis (ap- pendix 1; see the following discussion).
All character states were compiled into a total of 246 transformation series. In many cases, the states within a given transforma- tion series were so numerous and complex that it was more practical to handle them as a series of multiple binary transformations. Those cited transformation series that pre- sented interpretive problems (e.g., relative size, relative position) are discussed below. It should be noted that the sequence in which the character states are listed in a given trans- formation series does not imply transforma- tion additivity or polarity. Moreover, this phase of the analysis does not attempt to pos- it phylogenetic relationships: Outgroup com- parison, and subsequent hypotheses of char- acter polarity and directionality are addressed in the second phase (see Phylogenetic Anal- ysis below).
Dental nomenclature follows that of Rich (1981); refer to figure 2. In general, because the nomenclature is fairly consistent across the Erinaceomorpha, the majority of the lit- erature-based apomorphies are self-explana- tory (refer to fig. 2) and need no discussion. The characters pertaining to the molars are illustrated in figures 2a, b, an idealization of the occlusal surfaces of upper and lower tri- bosphenic molars (following Salay, 1969 and Rich, 1981). Stereo photographs of occlusal surfaces of representatives of each genus re- viewed in this analysis are presented in fig- ures 3-7.
Interpretation difficulties are almost exclu- sively confined to those transformation series that attempt to characterize size and shape in a nominal (i.e., noncontinuous) fashion. For example, “‘the hypocone is larger than the protocone’”’ (Storch and Qiu, 1991)—it is un- clear whether the size “‘larger”’ refers to the height of the cusps, or the gross size (vol- ume) of the cusps, or both. At first glance, this may seem trivial, but many fossil taxa, such as those that are represented only by dental material, are described and diagnosed based on such character states (appendix 1).
Herein I have tried to accurately define the size parameters to which I refer, however, there still remains the problem of visualizing size without the aid of controlled measure- ments (e.g., employing the use of calipers).
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i I ! |
encd end
Fig. 2. Occlusal view of idealized tribosphen- ic molars: (a) first upper molar; (b) first lower molar (taken from Rich, 1981). Abbreviations: cc = centrocrista (to include postparacrista and pre- metacrista); co = cristid obliqua; ecg = ectocin- gulum; ecgd = ectocingulid; efx = ectoflexus; encd = entocristid; enld = entoconulid; end = entoconid; hy = hypocone; hyd = hypoconid; hyld = hypoconulid; hyxd = hypoflexid; me = metacone; mec = metacrista (or postmetacrista); med = metaconid; meg = metacingulum; ms = mesostyle; msd = mesoconid; mt = metastyle; mtl = metaconule; pa = paracone; pac = para- crista (or preparacrista); pacd = postparacrista; pad = paraconid; pag = paracingulum; pcg = pre- cingulum; pmlc = premetaconule crista; pple = preparaconule crista; pprc = preprotocrista; pr = protocone; pred = protocristid; prd = protoconid; prl = paraconule; prgd = precingulid; ps = par- astyle; psc = postcrista; pscg = postcingulum; psgd = postcingulid; psmlc = postmetaconule crista; psplc = postparaconule crista; pspre = postprotocrista; st = stylocone; sts = stylar shelf; tb = trigon basin; tdb = trigonid basin; tlb = tal- onid basin; tlh = talonid notch; trn = trigonid notch.
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GOULD: DENTAL MORPHOLOGY OF HEDGEHOGS f
Fig. 3.
This difficulty became apparent on many oc- casions when I, along with W. R. Downs, reviewed the same specimen with consider- ably different impressions. Orientation is partly responsible for the variant interpreta- tions: When the tooth is in situ (.e., in the jaw), it does not sit on a level plane, there- fore the heights of the cusps appear different depending on whether they are viewed labi- ally or lingually. Because of the sheer mag- nitude of the number of specimens reviewed in this analysis (227), along with the varying size and fragility of the specimens, taking measurements for every cusp was precluded. Instead, the specimens (skull and jaws) were placed so that the occlusal surfaces were on approximately the same plane (a natural po- sition) and the relative heights of the cusps were recorded.
Other ambiguous characters concern cusp position illustrated, for example, in the state- ment “‘the paracone is lingual to the meta- cone’? (Koerner, 1940). As with size, de- scriptions of cusp position depend on which
Echinosorex gymnura (AMNH 115519) upper and lower jaw.
part of the cusp is referred to: the base or the apex. In many cases, especially the proto- cone and protoconid, the cusp is somewhat crescentic in shape (See figs. 3—7), and thus the apex extends lingually beyond the base of the cusp. The apex of the protocone, how- ever, becomes more aligned with its base with progressive wear (personal obs.). I have thus tried to standardize these relative-posi- tion characters by referring to only the base of the cusp. These revised positional defini- tions may not be in accord with the original intention of the author(s) who first observed and noted these characters (appendix 1), nev- ertheless, the base of the cusp is much less susceptible to wear, making its position less likely to be compromised.
Equally difficult to interpret is what con- stitutes a character or character state in the mind’s eye of another investigator. Is an enamel “*bead”’ on the labial side of the tooth equivalent to the presence of a labial cingu- lum? Or is a mediolateral crest extension of the protoconid on the p4 considered a distinct
AMERICAN MUSEUM NOVITATES NO. 3340
Fig. 4. Hylomys sinenesis (AMNH 10106) upper and lower jaw.
Fig. 5. Atelerix albiventris (AMNH 165804) upper and lower jaw.
2001 GOULD: DENTAL MORPHOLOGY OF HEDGEHOGS 9
Fig. 6. Erinaceus europeaus (AMNH 70611) upper and lower jaw.
Fig. 7. Hemiechinus auritus (AMNH 85309) upper and lower jaw.
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cusp (i.e., paraconid)? Fortunately, there are just a few examples where observer interpre- tation varies greatly. I have tried to adhere as closely as possible to the literal definitions of the characters reviewed (e.g., a labial enamel bead is not a cingulum, nor is a crest consid- ered a distinct cusp). In some cases, I found it necessary to introduce new transformation series to accommodate commonly expressed character states (e.g., cuspules) that could not be accommodated comfortably in an already cited transformation series. I did not include those anomalous character states (e.g., dou- ble apex on the p4 paraconule) that were un- likely to have any potential for phylogenetic inference, since they are all individually spe- cific, but it should be noted that such varia- tion occurs.
The most pervasive difficulty in the anal- ysis is apprehending the effects of wear on discrete dental characters. As discussed in the Introduction, wear is a consequence of many factors and processes, which ultimately results in the alteration of discrete dental characters at differing rates on an animal’s full complement of teeth, both deciduous and permanent. These ontogenetic differences may be easy to apprehend as the effects of wear in a large sample. In those cases where only a few specimens are readily available (or even exist), however, this type of ambi- guity could lead to unconscious character- state inference on the part of the investigator (Nixon, 1996).
Lastly, sample size itself may also pose problems: 25 individuals per taxon may not be a large enough sample to detect interspe- cific polymorphism, let alone the covariances with ontogenetic stages considered in the analysis.
SCORING OF CHARACTERS: All individuals (see appendix 2) included in this analysis were personally reviewed and appropriately scored for the listed transformation series (appendix 1). The maximum number of mul- tistates within a given transformation series is 5 G.e., O-4). Due to asymmetry, however, character coding is not as straightforward as 0—4. The fashion in which the asymmetrical data were recorded was designed to clearly indicate ‘“‘morphographic distribution”’ of the polymorphism in a single individual. For ex- ample, states in the left and right teeth de-
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scribed as [2,1] would mean the right tooth has state 2, and the left tooth has state 1. Because most statistical packages cannot ac- commodate entries with commas, necessitat- ing recoding for final analysis, coding of asymmetry starts with 5 and ends with 14. Coding is as follows:
(Oy Lor, [le 0] 5.4(O8 24 or 2, sO 6 [Os 3: or [3, 0] 7; [0, 4] or [4, 0] 8; [1, 2] or [2, 1] 9; [i 3]-6r [3,1] 10; Lt, 41 or (4, -1] 11; (2; 3] or [3, 2] 12; [2, 4] or [4, 2] 13; [3, 4] or [4, 3] 14.
Although many of these combinations of asymmetrical polymorphism do not exist in the taxon-specific matrices, it was more ef- ficient to recode all the possibilities through a linear editing function approach of the sta- tistical package in which the data were col- lected (see below) than to accomplish this task by hand.
A result of this coding method is the acute loss of asymmetry distribution (e.g., [1, 3] or [3, 1] 10). Because this analysis seeks only to acknowledge that dental asymmetry exists among erinaceids, without exploring its na- ture, the loss of distributional information on asymmetry is considered insignificant. The raw data are preserved in Gould (1997; ap- pendices 9-18).
As with virtually any other data set, miss- ing values are present. In those cases where observation of a character state was unequiv- ocally compromised by wear, the cell was left blank. Also, for those transformation se- ries that were not applicable to the taxon be- ing reviewed, the cells were also left blank. I did not code these data differently from other missing data, because operationally they are treated the same in a phylogenetic analysis that employs the parsimony princi- ple (Maddison, 1993; Nixon et al., 1994; Nixon, 1996).
TAXA REVIEWED: Of the 19 extant phylo- genetic species currently recognized (fig. 1b), 10 were considered in this analysis (appendix 2). Many of the living erinaceids are surpris- ingly rare in North American collections, which constrained the sampling criteria ac- cordingly. The optimal sampling parameters were as follows: (1) all specimens had to be accessible for personal review; (2) each tax- on had to be represented by an ideal of 25 individuals and a minimum of 10, and the
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sampling constrained, if possible, to one geo- graphic population; and finally, (3) each ge- nus had to be represented by at least 2 spe- cies. Criterion no. 2 may violate classical sampling criteria in that the sample size may be insufficient to clearly apprehend variation (Sokal and Rohlf, 1995), either due to a pau- city of specimens, or to a biased sampling within a population. Nevertheless, when dealing with the vertebrate fossil record, such parameters quickly become irrelevant in the face of inadequate sample sizes (e.g., one, or two specimens). Therefore, a small sample size of living taxa does not seem to be op- erationally any different than that of the fos- sil record, and in fact, a sample of 25 indi- viduals per taxon Is rare.
The limited sample size of this analysis also results from the need for personal re- view. As mentioned, North American collec- tions of erinaceids are limited, the majority of which are located at the Smithsonian Nat- ural History Museum and the American Mu- seum of Natural History. Thus, taxa chosen were predetermined by their availability in these two collections. Of the 7 recognized living genera (following Frost et al., 1991; fig. 1b), 5 were sampled. Due to the rarity of Podogymnura (Hylomyinae, 2 species rec- ognized) and Mesechinus dauuricus (Erina- ceinae, monospecific genus) in these collec- tions (fewer than 6 specimens per taxon were available), they were omitted from this anal- ysis. Echinosorex (fig. 3), currently consid- ered a monospecific taxon (E. gymnura) fol- lowing Corbet (1988) and Frost et al. (1991), is the only taxon that is represented by more than 25 individuals (32 were reviewed). This exception to the maximum sample size ex- ceeds the target sample size of 25 individuals from (presumably) one population from western Malaysia, and 5 individuals from the island of Borneo, which previously had been considered a separate (sub)species, E. gym- nura albus (Corbet, 1988). Given the avail- ability of these specimens and the nature of this project (inter- and intraspecific varia- tion), the addition of these specimens seemed appropriate.
The following genera were analyzed: Hy- lomys (fig. 4), Atelerix (fig. 5), and Erinaceus (fig. 6) each represented by 2 species; and Hemiechinus (fig. 7), represented by 3 spe-
GOULD: DENTAL MORPHOLOGY OF HEDGEHOGS sk
cies. Three hemiechinines were analyzed to ensure the inclusion of at least one taxon that was previously considered to be Paraechinus (Rich and Rich, 1971; Rich, 1981; Corbet, 1988; Frost et al., 1991). Twenty-five indi- viduals of the species Hylomys suillus, Eri- naceus amurensis, Atelerix algirus, and Hemiechinus hypomelas were not available for review, and therefore, smaller sample siz- es had to be accepted: 16, 11, 24, and 24, respectively.
OTHER DATA COLLECTED: Data regarding the relative age, sex, and geographic locali- ties of each individual (see appendix 2; see also Gould, 1997, appendices 9-18) have also been compiled; they constitute the var- iables against which discrete dental variation was tested for covariance.
The relative age categories—juvenile, ma- ture adult, worn teeth—are representative of wear stages, as there is no question that wear is principally a function of the age of an in- dividual (Brockie, 1959; Skoudlin, 1976, 1981; Gregory, 1976; Kahmann and Ves- manis, 1977; Vasilenko, 1988). The identifi- cation of juveniles is straightforward; it is based on the presence of deciduous teeth. The remaining two age categories are based on the following definitions: [category: worn teeth] those specimens identified as old run the gamut from having teeth worn to the roots to teeth worn just to the point where identification of certain discrete characters becomes murky (e.g., metaconule, cristae); [category: mature adult] all remaining indi- viduals that do not have deciduous teeth or morphology that is clearly compromised by wear. These categories may seem arbitrary or imprecise, but to estimate the age of a hedge- hog accurately is no simple task. Previous investigations regarding age estimation among erinaceines suggest that the only re- liable methods are: (1) measuring the relative dry weight of eye lenses, which increases with age (Morris, 1969, 1970, 1971); (2) not- ing the stage of epiphyseal fusion (Morris, 1971; Reeve, 1981; Dickman, 1988); (3) de- termining the number of periosteal growth lines in the lower jaw (Kristoffersson, 1971; Kratochvil, 1975; Dickman, 1988); and (4) observing dental wear stage (Brockie, 1959; Skoudlin, 1976, 1981; Gregory, 1976; Kah- mann and Vesmanis, 1977; Vasilenko, 1988).
12 AMERICAN MUSEUM NOVITATES
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TABLE 1 Results of Covariant Analyses
The percentages of polymorphic and asymmetric characters are calculated from the maximum of 246 transformation series minus those characters not applicable (Missing) to each taxon. The transformation series that covaried with sexual dimorphism, deciduous dentition, wear stage, and geographic locality are calculated based on the number of polymorphic transformation series per taxon.
Covariance Sexual Deciduous Wear Geographic Taxon (sample size) Missing Polymorphic Asymmetric dimorph. dentition stage locality Echinosorex gymnura (32) 6 103 (43.1%) 48 (20%) 0 0 3 (2.9%) 1 (0.04%) Hylomys sinensis (25) 20 TT (34.2%) 82 (36.4%) 0 0 0 0 Hylomys suillus (16) 18 67 (29.3%) 63 (28.0%) 0 1 (1.5%) 6 (8.9%) 0 Atelerix albiventris (25) 34 94 (44.5%) 84 (39.8%) 0 0 6 (6.4%) 0 Atelerix algirus (21) 28 80 (36.9%) 32 (14.7%) 0 0 4 (5.0%) 0 Erinaceus amurensis (11) 25 55 (25.0%) 9 (4.1%) 0 1 (1.8%) 6 (10.9%) 0 Erinaceus europaeus (24) 37 70 (33.7%) 60 (28.8%) 0 0 1 (1.4%) 0 Hemiechinus aethiopicus (25) 30 95 (44.2%) 66 (30.7%) 0 ¢) 3 (3.2%) 0 Hemiechinus auritus (25) 30 66 (30.7%) 27 (12.4%) 0 2 (3.0%) 6 (9.1%) 0 Hemiechinus hypomelas (19) 28 75 (34.6%) 48 (24.5%) 0 0 11 (14.7%) 0
Given the lack of access to fresh eye lenses and postcranial material for review of epiph- yseal fusion and of permission to take thin sections from hundreds of specimens for age determination, wear stage was deemed ac- ceptable for estimating age.
Admittedly, using a second age variable would have increased the rigor in this anal- ysis. However, an early analysis of cranial suture closure (basioccipital and premaxil- lary-maxillary-palatine) demonstrated that these sutures close at approximately the same time very early on in ontogeny (personal obs.; Gould, 1997), thus, they would not have provided any additional information re- garding the age of an individual.
The other variables—sex and geographic locality—were determined from specimen tags.
ANALYSES CONDUCTED: Data were initially collected in MicroSoft Excel 4.0 for the Macintosh. It was then subsequently trans- posed and imported into both StatView 4.1, and a promotional version of (SAS) JMP for the Macintosh. The vast majority of all of the discrete dental data analyses (DDA) were conducted using StatView 4.1. All taxon data matrices (Gould, 1997, appendices 9-18) were first reviewed for intraspecific variation, as well as asymmetry within a transformation series (DDA 1). The data were analyzed by generating frequency tables for each trans-
formation series across all the taxa (Gould, 1997, appendices 9-18). The results have been compiled in one table (appendix 3) for a global overview of variation. Identified in- terspecific polymorphism was then analyzed for covariance with sexual dimorphism (DDA 2), deciduous dentition and wear stage (DDA 3), and geographic locality (DDA 4). Bar chart cell plots were employed to visu- alize the distribution of the data, and their covariation with the variables noted (see ap- pendix 4 for examples).
RESULTS OF DISCRETE DENTAL ANALYSES
Discussion of the results of each analysis is as follows: only those transformation se- ries that decidedly covaried with the three variables considered—sex, relative age, and geographic locality—are herein discussed and illustrated. Table 1 is a compilation of the overall results of this analysis, and ap- pendix 3 is a comprehensive table of fre- quency of all the taxa and transformation se- ries that have been reviewed in this study.
Decidedly implies that the results were not equivocal. That is, the frequency distribu- tions did not require any ad hoc hypothesis to explain conflicting results. For example, in fig. 8a, the distribution of I2 posterior cus- pules is illustrated. In the juvenile, one con- dition is observed—present—whereas in the
2001
TS.7. 12, posterior cuspules:
aS vA A
me N
a. Juvenile
'TS.37 P3, posterior cingulum: 4.5
4 3.5 3 2.5 # 15
b.
Fig. 8.
Juvenile
Mature adult
Mature adult
An example of ambiguous distribution of character states; (a) I2 cuspules always present in
GOULD: DENTAL MORPHOLOGY OF HEDGEHOGS 13
(| 0 present BB 1 absent
Worn teeth
~ [ ] O present | iB 1 absent
Warn teeth
the juveniles and polymorphic for the adults (Hylomys suillus); (b) expression of the P3 posterior cingulum is a consequence of wear (Erinaceus amurensis).
other two age categories, both present and absent conditions are observed. It is unclear from this distribution whether we are looking at: (1) distinct deciduous morphology (pres- ence) and polymorphism in the adults; (2) the effects of wear in only some individuals; or (3) a poor sampling of juveniles resulting in no detection of polymorphism. These distri- butions become even more problematic when
it is unclear whether a juvenile’s teeth are deciduous or permanent. Consequently, all the taxon-specific character distributions that were ambiguous (like this one) were consid- ered equivocal.
In figure 8b, the distribution of the P3 pos- terior cingulum (present [0], absent [1]) strongly indicates that wear accounts for the observed polymorphism. Distributions such
14 AMERICAN MUSEUM NOVITATES
as this one were considered evidence of co- variation.
DDA 1: POLYMORPHISM AND ASYMMETRY: Polymorphism and asymmetry are prevalent in all of the taxa reviewed (table 1, appendix 3; see also Gould, 1997, appendices 9-18). For all the transformation series considered, overall polymorphism within a given taxon ranges from 25% to 44.5%, asymmetry being slightly more conservative, ranging from 4.1% to 39.8%.
The overall amount of polymorphism (and asymmetry) detected across and within 10 taxa and 246 transformation series does not seem too surprising considering the quantity of characters reviewed. What is surprising, however, are the characters that are consis- tently polymorphic across all the taxa—the number of upper canine roots, the number of P2 roots, the presence and absence of the P3 lingual lobe, and the shape of the P4 and condition of its lingual roots, to mention a few (appendix 3). These character states have all been cited in the literature as diagnostic for a taxon, either at the species level or higher (see appendix 1). Moreover, the poly- morphic presence and absence of an entire tooth (12, 13, Pl, and P3) within a species is even more disconcerting (appendix 1). With- out large series of individuals from a single population by which to detect such variation, these characters could be considered evi- dence of multiple species.
DDA 2: COVARIATION OF POLYMORPHISM AND SEXUAL DIMORPHISM: The results of this analysis suggest that there is no expression of sexual dimorphism in the discrete dental characters among the reviewed taxa, and most probably throughout the living erina- ceids as well.
DDA 3: COVARIATION OF POLYMORPHISM WITH RELATIVE AGE: All positively correlated results of polymorphism with the relative age of an individual (wear stage and deciduous vs. permanent teeth) are illustrated in table 2 (see also Gould, 1997, appendix 4).
In some taxa, the deciduous dentition can be quite different from the permanent teeth. Results of this analysis, however, indicate that among hedgehogs, the morphology of deciduous and permanent teeth is not easily distinguishable. Of the 10 taxa reviewed, only 3 distinctly demonstrate polymorphism
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in deciduous and adult teeth (see table 2): Hylomys suillus (upper canine size relative to postcanines); Atelerix algirus (presence of I3 posterior cingulum); and Hemiechinus auri- tus (P3 is reduced, dP3 protocone is present). (see also Gould, 1997, appendix 4, figs. 5, 18, and 34, respectively.)
Thirty-nine characters were found to be positively correlated with wear (table 2; see also Gould, 1997, appendix 4, figs. 2—4, 6— 17, 19-33, 36-51); those that are consistent- ly affected are: premolar cuspules, cingula, parastyle, and cristae. These wear-dependant characters are not tooth specific, they tend to be unfailingly distributed across almost all the teeth that exhibit that particular character state. For example, the parastyle is subject to wear on the P3, P4, M1 and M3; cingula are subject to wear on the upper canine through the M3, and the m2 (table 2). These wear patterns are directly correlated with occlusal surfaces of the parastyle: the P4 parastyle oc- cludes with the posterior crest on the lower canine; the M1 parastyle occludes with ml protoconule; the M2 parastyle occludes with m2 protoconule; and the M3 parastyle oc- cludes with the m3.
The wear of the cingula is not as clear cut. Only two of the four cingula (on a premolar, upper molar, or lower molar) are occlusal surfaces: the anterior and posterior cingula, which occlude with the protocones and pro- toconids, respectively. Wear of the labial and lingual cingula among erinaceids must be a result of diet (or “‘bug wear,’ sensu D. R. Frost). Hedgehogs have a varied diet, includ- ing: insects, snakes, eggs, small mammals, and small lizards (Lui, 1937; Krishna, 1956; Brockie, 1959; Burton, 1969; Herter, 1969; Campbell, 1973; Roberts, 1977; Merrit, 1981; Maheshwari, 1984; Corbet, 1988; also see Reeve, 1994 for a complete review), all of which can be abrasive to teeth. Hedgehogs are also known to dispatch relatively large invertebrates using their molars (Reeve, 1994; pers. obs.), instead of tearing with their incisors, or even canines. This observation would explain the wear of the labial cingu- lum recorded in this analysis.
DDA 4: COVARIATION OF POLYMORPHISM WITH GEOGRAPHIC LOCATION: Only one taxon, Echinosorex (fig. 3), exhibited dental varia- tion (presence/absence of I1) that conclusive-
2001 GOULD: DENTAL MORPHOLOGY OF HEDGEHOGS 15
TABLE 2 Transformation Series Found to Covary with Deciduous (=d) versus Permanent Dentition, Wear Stage (= w), and Geographic Variation (=g) Abbreviations follow those of Frost et al. (1991); ECHG = Echinosorex gymnura; HYLU = Aylomys suillus; ATXA = Atelerix albiventris; ATXG = A. algirus; ERIA = Erinaceus amurensis; ERIAE = E. europaeus; HEME = Hemiechinus aethiopicus; HEMA = H. auritus; HEMH = dH. hypomelas.
Taxon Transformation series ECHG HYLU ATXA ATXG ERIA' ERIAE HEME HEMA HEMH
Polymorphism: Deciduous ys. Permanent Dentition (total = 4) 16. 13, posterior cingulum d 18. UC, size relative to postcanines d 39. P3, morphology 52. P3, roots
af
Polymorphism: Wear Stage (total = 39) 21. Upper canine, anterior cingulum WwW 23. Upper canine, posterior cuspule Ww 30. P2, posterior cuspule Ww 37. P3, posterior cingulum Ww 41. P3, protocone position relative to paracone w 48, P3, parastyle Ww 54, P4, hypocone w 61. P4, protocone-hypocone crest w w 62. P4, metastyle Ww 63, P4, parastyle Ww 64. P4, anterior cingulum w 73, M1, metaconule w 74. M1, metaconule shape Ww 75. M1, postmetaconule crista Ww 76. M1, protocone height w 85. M1, centrocrista 89. M1, paraconule 90. M1, paraconule crista W 98. M1, parastyle w 100. M1, cingulum w 101. M1, lingual cingulum w w 104. M1, labial cingulum w 114. M1, paraconule Ww 125. M2, anterior cingulum Ww W 126. M2, labial cingulum Ww 127. M2, posterior cingulum 136. M3, parastyle w 137. M3, anterior cingulum w 139. M3, posterolabial cingulum 154. LC, position relative to preceding incisor 155. LC, anterior midline crest 157. LC, posterior ridge Ww 166. p2, cingulum w 169. p2, anterior midline crest Ww 214. ml, hypocristid Ww Ww 217. mt, entoconulid w 227. m2, paraconid swelling Ww 231. m2, posterior lingual extension Ww 242. m2, labial cingulum Ww
2 = = =
z=
z=
Polymorphism: Geographic Variation (total = 1) 4. Tl, presence/absence g
16 AMERICAN MUSEUM NOVITATES
ly covaried with geographic locality (table 2; see also Gould, 1997, appendix 4, fig. 1).
Echinosorex is the largest member of the Erinaceidae, and as well, the largest of the living lipotyphlans (Frost et al., 1991). Its known distribution extends throughout the Indonesian Peninsula and the Malayan Ar- chipelago (Lim, 1967), to include the islands of Burma, Sumatra, Malaya, Thailand, and Borneo. The genus Echinosorex has been previously thought to contain at least three [sub]species: FE. dealabatus, E. alba, and E. gymnura (Corbet, 1988). I bracket the [sub] as these taxonomic designations have not been consistent. Recent revisions of the tax- onomy of Erinaceidae considered Echinoso- rex to be a monospecific taxon (Corbet, 1988; Frost et al., 1991). In this analysis, the absence of the Il seems to be apomorphic for the population in Borneo, although this is based on a review of only five specimens.
BRIEF SYNopSIS: Polymorphism and asym- metry were discovered to be quite common across all of the taxa reviewed in this anal- ysis. Of the 246 transformation series con- sidered, 204 (83%) were found to exhibit in- traspecific variation. Of all the polymor- phism exhibited, very little could be attri- buted unequivocally to any of the variables (age, sex, geography) considered in this anal- ysis. This does not suggest that the discrete dental characters do not covary with these variables (except perhaps for sexual dimor- phism); it simply suggests that it is very dif- ficult to discern covariation from random in- dividual variation.
Aylomys suillus (fig. 4) and Erinaceus amurensis (fig. 6) have the least amount of polymorphism (and asymmetry) relative to all of the taxa reviewed, with Afelerix albi- ventris (fig. 5) exhibiting the most. The rel- atively low frequencies of polymorphism in the two above mentioned taxa may be attrib- utable to small sample sizes: 16 and 11 in- dividuals, respectively.
PHYLOGENETIC ANALYSIS
METHODS AND JUSTIFICATION
The results of the discrete dental analyses of the 246 transformation series reviewed, in- dicated that this character suite exhibits con- siderable amounts of variation across all tax-
NO. 3340
onomic levels within the Family Erinaceidae. Currently, there is no consensus on the treat- ment of variable or polymorphic characters in a phylogenetic analysis (see Weins 1998, and Kornet and Turner, 1999 for a compre- hensive review of methods). Nor is there a consensus on whether or not they should even be included in a phylogenetic analysis (Nixon and Wheeler, 1990; Nixon and Davis, 1991; Kornet and Turner, 1999) despite em- pirical data to the contrary (Campbell and Frost, 1993; Nixon and Carpenter 1993; Nix- on et al, 1994; Weins, 1995, 1998).
It is not my intention that this analysis test methods of phylogenetic reconstruction, or even reconstruct a phylogeny of the Erina- ceidae. My intention is to inquire only into the phylogenetic resolving power of discrete dental characters in the absence of all other data. Therefore, given that 83% of the dis- crete dental characters exhibited intra- and/ or interspecific variation, I constrained the phylogenetic analysis to best maximize the resolving power of the “‘fixed’’ characters. The question of the phylogenetic resoloving power of polymorphic characters within the Erinaceidae will have to wait for future study.
Interspecific variation was set to a maxi- mum number of three species for a given transformation series. That is, if three or more species demonstrated considerable in- terspecific variation for a given transforma- tion series, I omitted it from the analysis. In sum, 100 transformation series were retained (appendix 4).
Recorded variation in three of the trans- formation series included in this analysis— Il presence/absence, P3 morphology, and P4 hypocone—exhibit positive covariation with geographic locality, deciduous dentition, and/ or wear stage, respectively (table 2). It should be noted that I included a transfor- mation series that is known to be affected by wear for two reasons: (1) it was found to covary with wear in only one taxon, Hem- iechinus hypomelas, which can easily be ac- counted for a posteriori to any analysis; and (2) the presence/absence of the P4 hypocone has historically been considered apomorphic at some taxonomic level within the Erina- ceidae (Butler, 1948, 1988; Novacek, 1985, 1986; Frost et al., 1991).
2001
The data were analyzed following the cri- teria set by Gould (1995) and Frost et al., (1991) for outgroups (see below) and anal- ysis parameters (e.g., PAUP, branch swap- ping methods). These criteria were rigorous- ly adhered to in order to maximize the com- parability of the discovered trees. Outgroups employed are the tenrecoids and soricoids. The fossil taxon, leptictids, was omitted from this analysis. Omission of this taxon does not affect the topology of the trees of either Frost et al. (1991) or Gould (1995), thus its inclu- sion did not seem pertinent.
I have coded the outgroups for as many of the transformation series for which I felt comfortable in making statements of “‘pri- mary” homology (di Pinna, 1991). The sometimes extreme differences in dental morphology (i.e., tribospheny vs dilamdo- donty and zalamdodonty) among the ingroup (erinaceids) and outgroups (soricoids and tenrecoids) prohibits statements of homolo-
All characters were polarized according to the outgroup criterion (see Nixon and Car- penter, 1993), and all multistate transforma- tion series were left unordered. Although I am not comfortable leaving the multistates unordered, many of the position or size-re- lated characters lack evidence to justify ad- ditivity (e.g., entoconid size: (0) > hypocon- id; (1) > paraconid; (2) = to both cusps; (3) > both cusps).
In order to test the phylogenetic resolving power of any data set, in this case discrete dental characters, a standard must be used against which to test it. As mentioned in the Introduction, Frost et al. (1991) and Gould (1995) posited hypotheses of the erinaceid phylogenetic relationships based on general morphology. These hypotheses are congru- ent, despite the somewhat different data sets analyzed (both in terms of taxonomic and character composition, see fig. 1) and are thus employed as the standard with which to compare the results of Data Set 1.
As a secondary internal test, Gould’s den- tal data (1995; Data Set 2 = 29 characters) were isolated and reanalyzed. Phylogenetic analysis 2a includes only those 10 taxa re- viewed in this analysis. PA 2b considers the 19 living taxa included in Gould’s original analysis, as well as that of Frost et al. (1991).
GOULD: DENTAL MORPHOLOGY OF HEDGEHOGS As?
As with Data Set 1, the same outgroup cri- teria were employed.
Given that this analysis does not set out to reconstruct phylogenetic relationships, but rather to look at the topological effects of using a single suite of characters for phylo- genetic inference within hedgehogs, incon- gruence length difference and significance tests (Mickevitch and Farris, 1981) were not considered here.
The computer-assisted parsimony program PAUP (Swofford, 1993) was used to analyze the data. A heuristic search was conducted, using random tree stepwise addition, and tree bisection branch-swapping algorithms. The outgroup option was employed, and both ACCTRAN and DELTRAN optimizations were considered.
Abbreviations: CI = consistency index; RI = retention index; RC = rescale consistency index.
RESULTS OF PHYLOGENETIC ANALYSIS
PHYLOGENETIC ANALYSIS 1: DATA SET 1: Analysis of the 100 transformation series from the discrete dental analysis and 10 taxa discovered six trees of 105 steps, with the following statistics (excluding uninformative characters): CI = 0.634; RI = 0.528; and RC = 0.357; the strict consensus and the Adams tree are depicted in figs. 9a and b, respec- tively. In all the trees discovered, every poly- typic genus is rendered paraphyletic (except Erinaceus), and the monophyly of both ex- tant subfamilies is challenged (compare with figs. la and b).
PHYLOGENETIC ANALYSIS 2: DATA SET 2: Analysis of Frost et al.’s (1991) dental data across the 10 taxa reviewed in this investi- gation discovered 4 trees: length 38; CI = 0.816, and 0.80 (excluding uninformative characters); RI = 0.897; RC = 0.732. The strict consensus tree and the Adams tree are the same (fig. 9c). The only genus discovered to be monophyletic is Erinaceus, both sub- families are rendered paraphyletic.
Analysis of Frost et al.’s (1991) dental data and the 19 living taxa they considered dis- covered 9 trees: length: 41; CI = 0.756 and 0.744 (excluding uninformative characters); RI = 0.917; RC = 0.694. The strict consen- sus and Adams tree are illustrated in figs. 9d
18 AMERICAN MUSEUM NOVITATES
soricoids
tenrecoids
Echinosorex gymnurus FAylomys suillus Hemiechinus aethiopricus Alelerix algirus Hemiechinus auritus Frinaceus europeaus Alelerix albiventris Hemiechinus hypomelas Hylomys sinensis
b.
a. Frinaceus amurensis
Fig, 9;
soricoids
tenrecoids
Echinosorex gymnurus Ayloniys suillus Hemiechinus dethiopricus Atelerix algirus Hemiechinus auritus Erinaceus europeaus Alelerix albiventris Hemiechinus hypomelas Hylomys sinensis
Erinaceus amurensis Cc.
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soricoids
tenrecoids
Echinosorex gymnura Hylomys suillus Hylomvys sinensis Hemiechinus aethiopicus Atelerix albiventris Atelerix algirus Hemiechinus hypomelas Hemiechinus auritus Frinaceus europeaus
Erinaceus amiurensis
(a) Phylogenetic analysis 1 (data set A); strict consensus tree; (b) majority rule tree; (c)
phylogenetic analysis 2a (Gould, 1995) strict consensus tree. (d) Phylogenetic analysis 2b (Gould, 1995)
strict consensus tree; (e) majority rule tree.
and e, respectively: hylomyine monophyly is challenged; Hylomys is never discovered to be a member of that group. Moreover, mono- phyly of all the living genera is suspect ex- cept for Erinaceus.
BRIEF SYNopsiIs: The recovered trees for all three analyses differ in overall topology both among themselves and with the hypotheses posited by Frost et al., (1991) and Gould (1995; figs. la and b). In each of the discov- ered topologies, all the taxa were rendered paraphyletic. Interestingly, the most parsi- monious trees discovered in all three analy- ses were seemingly well supported, as evi- denced by the high indices.
The apomorphy lists for two discovered trees are presented in appendices 5 and 6 with their respective data matrices. In all three analyses, Tree #1 was selected as the token topology from which to generate an apomorphy list (the strict consensus trees for the analyses are depicted in figs. 9a, c, and d). This arbitrary decision was based on the fact that not one of the discovered trees re- motely approximates any of the previously posited phylogenetic hypotheses that are based on all available morphological data (see Corbet, 1988; Frost et al., 1991; Gould, 1995).
The purpose of this analysis is not to pro- pose a phylogenetic hypothesis, but to ex- plore the phylogenetic resolving power of the discrete dental characters. Given the incon- gruous results with the most recent hypoth- eses of extant erinaceid relationships (Corbet, 1988; Frost et al., 1991; Gould, 1995), de-
tailed discussion of character support is fore- gone.
DISCUSSION
Briefly, the results of this investigation are: (1) variation is discovered to be rampant both inter- and intraspecifically, as well as within an individual; (2) correlation of some polymorphic characters with wear stage is demonstrated, although it is not consistent across the taxa reviewed; (3) polymorphism as a result of morphological difference be- tween deciduous and permanent dentition is discovered to be minimal and very difficult to detect without large sample sizes; (4) clin- al variation and sexual dimorphism of dis- crete dental characters are rare or nonexistent (respectively) among the taxa reviewed; and (5) dental characters, as a partitioned data set, recovered estimates of phylogeny that are globally incongruent with those based on comprehensive morphological data sets.
These results are not surprising. Variation of discrete dental characters across many mammalian taxa is already well documented (see Introduction). Within the Erinaceidae, it seems that the magnitude of discrete dental characters cited in the literature is a result of oversplitting of character transformations.
Wear is the primary cause of the altering of appearance of specific dental characters. It is not exclusively a function of age, but may also be a consequence of geographically (or individually) varying diets and/or individual pathology. For example, some individuals of
2001
soricoids tenrecoids Echinosorex gymmura Pedogymnura aureospinula Pedogymnura truei Hylomrys suillus Hylomys sinensis sabes hainanensis emiechinus aethiopicus Hemiechinus micropus Atelerix sclateri Alélerix albiventris Atelerix algirus Alelerix frontalis Hemiechinus hypomelas Mesechinus dauuricus Hemiechinus collaris Hemiechinus auritus Erinaceus concolor Frinaceus europeaus d. Erinaceus amurensis
Fig. 9. Continued.
Hemiechinus auritus were observed to have filled open cavities with sand grains (person- al obs.), a condition most likely due to ser- endipity. Nevertheless, a desert-dwelling hedgehog inadvertently ingesting sand parti- cles during a meal may not only fill a cavity, it is most likely going to wear down its teeth at a much more rapid rate (and in a different fashion) than a hedgehog living on a British Isle that takes in fine dirt and debris with its diet of earthworms and insect larvae (see Reeve, 1994, for a comprehensive review).
Theoretically these wear patterns could be apomorphic at some taxonomic level: it has been demonstrated that wear patterns can be indicative of behavioral characteristics: e.g., grazer vs. browser, habitat conditions, and even preferred diet (Solounias and Dawson- Saunder, 1988; Hayek et al., 1991; Solounias and Moelleken, 1992a, b, 1993; Solounias and Hayek, 1993). Hedgehogs are opportu- nistic feeders, the only constraint on their diet being environment; therefore, such wear patterns cannot be used as statements of ho- mology. In this analysis, I attempted to tease out ontogenetic variation (e.g., a function of wear) from ontological variation (e.g., sexual dimorphism, clinal or individual variation) which proved to be very difficult.
Although it is clear that wear occurs and that it alters tooth morphology over time, it is not easy to demonstrate empirically that wear is the principal cause of much of the observed variation. This is evidenced by the fact that 204 of the 246 characters reviewed were discovered to vary intraspecifically,
GOULD: DENTAL MORPHOLOGY OF HEDGEHOGS 19
soricoids
tenrecoids
Echinosorex gymnura Podogymmura truei Podogymnura aureospinula Hylomys suillus Aylomys sinensis Hylomys hainanensis Hemiechinus aethiopicus Hemiechinus micropus Alelerix sclateri
Atelerix albiventris Alelerix algirus
Atelerix frontalis Hemiechinus hypomelas Mesechinus dauuricus Hemiechinus collaris Hemiechinus auritus Erinaceus concolor Erinaceus europeaus
e. Erinaceus amurensis
and of these, only 46 (see table 2 and appen- dix 4) could unequivocally be attributed to wear. Clearly, many more of the polymor- phisms recorded in this analysis are a direct result of wear early in ontogeny; however, in most cases, little or no evidence of mechan- ical wear can be observed with a standard microscope. Only a comprehensive review of the various stages of molar eruption could demonstrate that the cingula were being worn off very early in the animal’s life. Without adequate sample sizes (and in some cases sophisticated methods of visualization [e.g., SEM scans]), the subtle topological manifestations of wear are not apprehendable using standard multivariate statistics.
To muddy the waters even more, premo- lars are both deciduous and permanent. In some mammalian taxa, deciduous dentition is different from that of the adult dentition; among erinaceids, however, deciduous and permanent dentition are quite similar (Kin- dahl, 1959)—only 4 characters across 3 taxa exhibit differing morphology (table 2). Nev- ertheless, I suspect that the low frequency of polymorphism due to distinct deciduous and permanent dentition is subsumed in the var- iation of the adult dentition and/or the reten- tion of milk teeth into adult life. With respect to the latter, without either clear signs of tooth eruption or X-rays, the nature of the variation is ambiguous. To compound the problem, deciduous teeth, like adult denti- tion, most likely vary intraspecifically, as well through wear.
Clinal variation and sexual dimorphism,
20 AMERICAN MUSEUM NOVITATES
expressed in the dental morphology, are even more elusive, if they exist at all. Among the sampled taxa, sexual dimorphism was found to play no role in the polymorphism. I had not anticipated that any teeth, other than pos- sibly the canines, would exhibit secondary sexual characteristics. Sexual dimorphism has never been demonstrated within the ex- tant hedgehogs; however among the fossil taxa, it has been suggested that the giant Ital- ian Miocene hedgehog, Deinogalerix ex- pressed sexual dimorphism in its overall size and number of premolars (Freudenthal, 1972; however see Butler 1980). Personal obser- vation of series of Echinosorex gymnura in- dicated (to me) that this taxon may also ex- press sexual dimorphism in the size of the skull.
Although it was demonstrated that there is some geographic variation in discrete dental data (i.e., Echinosorex), it should be noted that only one character (#4; presence/absence of I1) of the 246 reviewed, across 10 taxa could be directly correlated with geographic location: Echinosorex gymnura, a monotypic taxon distributed both on the mainland of In- dochina and the Indonesian and Malayan is- lands, exhibits geographically delimitable variation (see DDA 4, Results). This varia- tion may be in fact apomorphies indicating more than one phylogenetic species, not clin- al variation. This hypothesis has not been rigorously tested, and without larger sample sizes, I decline to re-establish another species of Echinosorex.
Results of the phylogenetic analyses of three overlapping discrete dental data sets in- defatigably indicate that discrete dental char- acters, in the absence of all other morpho- logical data, are insufficient for addressing questions of historical relationships among the extant taxa reviewed. This is evidenced by the fact that all of the discovered trees (Data set 1 and 2a/b) posit paraphyly and/or polyphyly of all the extant groups, across all taxonomic levels (see Butler, 1948; Rich, 1981; Corbet, 1988; Frost et al. 1991; Gould, 1995).
Given the frequency of polymorphism dis- covered in the discrete dental analysis, these results are not terribly surprising. What is disturbing is the number of trees discovered in each analysis and their respective indices.
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The maximum number of trees discovered for all of the data sets was 9, and the lowest CI was 0.63. If there were no other previ- ously postulated hypotheses of relationships that strongly corroborated one another, these hypotheses of phylogenetic relationships, based strictly on the number of discovered trees and their strong stem support, would be considered robust. One must not summarily dismiss these results as coincidental. We know that a mammal tooth is specialized based on its ability to occlude with its coun- terpart. One would assume, that the variation would be somewhat consistent to ensure that the teeth still remain functional (i.e., oc- clude), and further, that wear would perhaps enhance occlusion, consequently making many of these discrete dental characters de- pendent on one another (evocative of con- certed evolution and/or concerted ontogeny).
The principal questions being addressed in this investigation relate to how reliable den- tal data are as the sole source of phylogenetic inference for the fossil record. Given the re- sults of the discrete dental analysis, the caus- es of dental variation are elusive. I suspect that many of the variables reviewed in this analysis play some role in the exhibited var- iation. However, current methodology may be inadequate for teasing out which morpho- logic variation is real and which is an artifact of wear. Adding to these doubts are the well- supported results of the phylogenetic analy- ses that hypothesize nonindependence of characters and global paraphyly among the extant taxa of hedgehogs. In light of these results, I would be reluctant to place much weight on the phylogenetic resolving power of this particular suite of characters in the absence of other data. More specifically, I would hesitate to propose a taxonomy of fos- sil erinaceids based on fragmentary jaws and isolated teeth.
Within the field of paleomammalogy, the reliance on dental morphology as the sole in- dicator of phylogenetic affinities is fairly common. Understandably, this reliance is in direct response to what most researchers studying fossil mammals (especially small mammals) have to work with—isolated teeth or fragmentary jaws. Enamel survives oth- erwise harsh deteriorative and/or erosive en- vironments. Among some groups of mam-
2001
mals (e.g., dryolestids, triconodonts, ptilo- donts, and taeniolabids), teeth and jaws are the only record we have indicating that a lin- eage once existed. As a result, mammalian paleontological literature is replete with de- velopmental odontology, discrete dental mor- phology, microwear, and odontological mor- phometrics as standard methods for deter- mining phylogenetic relationships among certain taxa. If the dental data across all mammalian taxa is similar in behavior to that discovered within the extant Erinaceidae, these data may be misleading us. Certainly, it would be faulty reasoning to presume a priori that this dental homoplasy phenome- non is global for the Mammalia; neverthe- less, it casts doubt on the reliability of such data, especially when a cursory review of the literature suggests that similar observations are common within other mammalian taxa (see Introduction), to include Homo sapiens (Hillson, 1986; Melvin Moss, personal com- mun.).
Given the results of this analysis and oth- ers, it seems wise that, when possible, mea- sures should be taken to test the phylogenetic signals of the dental data on living taxa be- fore applying them to the fossil record. Nor should this type of approach be exclusive to mammalian teeth—all seriously depauperate data sets (those that use only one particular system of the animal to reconstruct evolu- tionary histories, should be rigorously tested before weighting them a priori (see Naylor and Marcus, 1994, and Sanchez-Villagra and Williams, 1998, for other methods of testing such data for application to the fossil record).
CONCLUSION
The results of the analyses of the discrete dental data conducted in this investigation strongly indicate that the expression of many characters commonly used (i.e., parastyle, cingula, cristae) to diagnose fossil erinaceid taxa are compromised by wear early in on- togeny (and in many cases little or no evi- dence of mechanical wear can be observed); they are subject to intractable, and global in- tra- and interspecific variation, and/or they are subject to concerted evolution. These data suggest further that intraspecific varia- tion, not unexpectedly, increases with sample
GOULD: DENTAL MORPHOLOGY OF HEDGEHOGS a
size (discrete dental analysis and phyloge- netic analysis). This issue is most pertinent within the discipline of paleontological sys- tematics. Not only is there scant material for review; in more instances than not, the taxon under consideration has no close living rep- resentatives (e.g., sauropods, parieasaurs, or nectridians) from which to get a better un- derstanding of the nature of the available fos- sil material. In such cases, there are no al- ternatives but to use the available material— a poor estimate of phylogenetic relationships may be preferable to no estimate of relation- ships at all.
For those taxonomic groups that have both living and extant representatives, a rigorous investigation of the phylogenetic signal of the available data for incomplete fossil taxa should be a prerequisite to any phylogenetic reconstruction (see Naylor and Marcus, 1994). As with any other data considered, such an investigation would minimally sat- isfy some of the criteria of a more rigorous methodological approach for phylogenetic inference by identifying characters too plas- tic to be useful (see Nixon and Davis, 1991, for an overview of the problems).
In sum, the factors reviewed herein—wear stage, clinal variation, gratuitous variation, and nonindependence of characters—can greatly alter our interpretation of the fossil record when the only evidence being re- viewed consists of teeth. Without consider- ation of these problems, the fossil species di- agnosed on such data must consequently af- fect all hypotheses of speciation events, mi- grations patterns, and hypotheses of evolutionary processes.
The results of this analysis presented here pertain only to the taxa that have been re- viewed herein. Admittedly it is difficult, if not impossible, to demonstrate empirically that these results also pertain to the erinaceid fossil record. Nevertheless, the usefulness of dental data for reconstructing their phyloge- netic histories is now undeniably suspect.
ACKNOWLEDGMENTS
I thank the following individuals for their invaluable contributions to this investigation: William R. Downs for helping me to collect much of the discrete dental data and for his
22 AMERICAN MUSEUM NOVITATES
willingness to discuss many of the issues pre- sented herein; Leslie E Marcus for his guid- ance in the statistical analyses conducted in this investigation; and J. Peter Meyer, for helping me compile some of these data and for his support throughout the duration of this project.
Without the aid and support of the Divi- sion of Vertebrate Paleontology at the Amer- ican Museum of Natural History (AMNH), this work would not have been possible. I thank the following funding agencies for fa- cilitating this study: Smithsonian Collections Study Grant, and the Theodore Roosevelt Memorial Fund, sponsored at the AMNH. I would also like to thank the departments of mammalogy at these institutions for making their collections available for study. I specif- ically want to thank Linda Gordon at the Smithsonian Institution who was most help- ful during the many months of data collec- tion in the Mammal Division.
Much appreciation goes to Marilyn Fox for molding and casting the extant teeth, Lor- raine Meeker for photographing and mount- ing them, and to Tom W. French for contrib- uting to some of the literature reviewed.
Thanks go to Darrel R. Frost, Leslie FE Marcus, Malcolm C. McKenna, Marc A. Carrasco, John Wible, and John Hunter for their comments on early drafts of this man- uscript.
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Determination of dietary adaptations of extinct ruminants through premaxillary analysis. Lethaia 26: 261-268. Solounias, N., and L-A. Hayek
1981.
1993.
1993. New methods of toothwear analysis and application to dietary determina- tion of two extinct antelopes. J. Zool. London 229: 421-445.
Stevens, M. S. 1977. Further study of Castolon local fauna
(Early Miocene) Big Bend National
2001
Park, Texas. Pearce-Sellards Ser. Texas Mem. Mus. 28: 1—69. Storch, G., and Z. D. Qiu 1991. Insectivores (Mammalia: Erinaceidae, Soricidae, Talpidae) from Lufeng hom- inoid locality, Late Miocene of China. Geobios (24) 5: 601-621. Surin, V. L., Bannikova, A. A., Tagiev, A. E, Oso- kina, A. V., and N. A. Formozov 1997. Molecular taxonomy of hedgehogs (Er- inaceidae, Insectivora) of Northeastern Paleartic: testing a new method. Dokl. Biol. Sci. 353: 156-158. Swofford, D. L.
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insectivore-primate transition. Bull. Am. Mus. Nat. Hist. 140: 17-57. Van Valen, L. 1967. New Paleocene insectivores and insec- tivore classification. Bull. Am. Mus. Nat. Hist. 135: 217-284. 1962. A study of fluctuating assymetry. Evo- lution 16: 125-142. Vasilenko, V. N. 1988. Age and sex structure in the white- chested hedgehog Erinaceus concolor (Martin) from the Caucasus. Ekologiya (Sverdlovsk) 19: 45—49. English trans- lation in Sov. J. Ecol. July—August 1988: 220-223. Viret, J. } 1938. Etude sur quelques Erinaceides fossiles specialement sur le genre Paleoerina-
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28 AMERICAN MUSEUM NOVITATES NO. 3340 APPENDIX 1 Transformation Series Considered in Discrete Dental Analyses Numbering starts at 4 to maintain consistency with the numbering in each of the taxon matrices in Gould’s dissertation (1997 [1—3 are specimen number, sex, and age, respectively]). 4, I]: (Q) present; (1) absent. 33. P3, posterolingual cingulum: (0) present; (1) absent
10. {1.
12.
13.
14.
15.
16.
17. 18.
19.
20.
21.
pies
23.
24.
25. 26.
Pa 28. 29. 30. 31.
aah
Il, size: (0) normal; (1) enlarged (Rich, 1981; Butler, 1948; Frost et al., 1991; Gould, 1995).
I2: (0} present; (1) absent.
12, posterior cuspule: (0) distinct; (1) weak; (2) absent. 12, size relative to I3: (0) greater; (1) approximately equal; (2) smaller (Novacek, 1985, 1986; Frost et al., 1991; Gould, 1995).
12, position: (0) displaced medially; (1) not (Rich, 1981; Frost et al., 1991; Gould, 1995).
13: (0) present; (1) absent (Rich, 1981).
I3 roots: (0) one; (1) two separate; (2) two fused (Mat- thew, 1929; Butler, 1948; Rich, 1981; Robbins and Setzer, 1985; Corbet, 1988; Frost et al., 1991; Gould, 1995).
13, paracone position: (0) directly ventral to the ante- nor root; (1) not (Rich, 1981).
I3, shape: (Q) nearly rectangle in occlusal view; (1) not (Rich, 1981).
I3, paracrista: (O) well developed; (1) not (Matthew, 1929; Rich, 1981).
I3, metacrista: (0) well developed; (1) not (Matthew, 1929; Rich, 1981).
13, posterior cingulum: (0) well developed; (1) not (Rich, 1981).
I3, posterior cuspule: (0) distinct; (1) weak; (2) absent. Upper canine size relative to postcanines: (0) signifi- cantly larger; (1) slightly larger; (2) approximately equal. Upper canine, roots: (0) two; (1) one; (2) two fused (But- ler, 1948; Rich, 1981; Robbins and Setzer, 1985; No- vacek, 1985; Novacek et al., 1985; Corbet, 1988: Frost et al., 1991; Gould, 1995).
Upper canine size relative to I3: (0) greater; (1) approxi- mately equal; (2) smaller (Matthew, 1929; Butler, 1948; Rich, 1981; Corbet, 1988; Frost et al., 1991; Gould, 1995). Upper canine, anterior cingulum: (0) present; (1) absent (Matthew, 1929; Rich, 1961).
Upper canine, posterior cingulum: (0) present; (1) absent (Matthew, 1929; Rich, 1981).
Upper canine, posterior cuspule: (0) distinct; (1) weak; (2) absent.
P1: (0) present; (1) absent (Butler, 1948; Frost et al., 1991; Gould, 1995).
Pl, anterior cuspule: (0) distinct; (1) weak; (2) absent. P1, posterior cuspule: (0) distinct; (1) weak; (2) absent (Butler, 1948).
P1, roots: (0) one; (1) two.
P2: (0) present; (1) absent.
P2, anterior cuspule: (0) distinct; (1) weak; (2) absent. P2, posterior cuspule: (0) distinct; (1) weak; (2) absent. P2, roots: (0) two separate; (1) one; (2) two fused; (3) three (Butler, 1948; Frost et al., 1991; Gould, 1995). P3: (0) present; (1) absent.
34,
35.
36.
37,
38.
39,
40.
4l.
42.
43.
44,
45. 46.
47.
48.
49. 50.
51.
Se
53.
54.
55.
56.
(Munthe and West, 1980).
P3, lingual lobe: (0) present; (1) vestigial or absent (But- ler, 1948; Rich, 1981; Corbet, 1988; Frost et al., 199t- Gould, 1995).
P3, lingual cingulum: (0) strong; (1) weak; (2) absent (Munthe and West, 1980; Rich, 1981).
P3, labial cingulum: (0) present; (1) absent (Munthe and West, 1980; Rich, 1981).
P3, posterior cingulum: (0) present; (1) absent (Rich, 1981).
P3, posterior cingulum condition: (0) small; (1) large (Rich, 1981).
P3: (0) normal; (1} reduced (Butler, 1948; Stevens, 1977; Munthe and West, 1980; Rich, 1981; Robbins and Set- zer, 1985; Novacek, 1985; Novacek et al., 1985; Corbet, 1988; Frost et al., 1991; Gould, 1995).
P3, protocone: (0) present; (1) absent (Butler, 1948, Stevens, 1977; Munthe and West, 1980; Rich, 1981-; Robbins and Setzer, 1985; Novacek, 1985; Novacek et al., 1985; Corbet, 1988; Frost et al., 1991; Gould, 1995), P3, protocone position relative to the paracone: (0) ante- rior; (1) posterior; (2) adjacent (Butler, 1948).
P3, protocone height relative to the paracone: (0) approx- imately twice as small; (1) much smaller (Rich, 1981).
P3, paracone shape: (0) conical; (1) crescentic (Munthe and West, 1980).
P3, postparacrista: (0) present; (1) absent (Rich, 1981; Munthe and West, 1980).
P3, metacone: (0) present; (1) absent.
P3, centrocrista: (0) present; (1) absent (Munthe and West, 1980).
P3, hypocone: (0) present; (1) vestigial or absent (Munthe and West, 1980; Butler, 1948; Gould, 1995). P3, parastyle: (0) strong; (1) weak; (2) absent (Stevens, 1977).
P3, metastyle: (0) present; (1) absent (Rich, 1981).
P3, preparacrista: (0) present; (1) absent (Stevens, 1977; Rich, 1981).
P3, preparacrista extension: (0) to parastyle; (1) not (Stevens, 1977; Rich, 1981).
P3, roots: (0) three; (1) fewer (Butler, 1948; Rich, 1981; Corbet, 1988; Frost et al., 1991; Gould, 1995).
P4, shape: (0) quadrate; (1) rectangular; (2) triangular (Crusafont et al., 1955; Black et al., 1980).
P4, hypocone: (0) present; (1) absent (Butler, 1948, 1988, Novacek, 1985; Novacek et al., 1985; Frost et al., 1991). P4, hypocone height relative to the protocone: (0) small- er; (1) approximately equal (Matthew, 1929; Black et al., 1980).
P4, hypocone gross size relative to protocone: (0) small- er, (1) approximately equal (Rich, 1981; Storch and Qiu, 1991).
2001 GOULD: DENTAL MORPHOLOGY OF HEDGEHOGS APPENDIX 1 Continued
57. P4, protocone position with respect to that of the para- 79. M1 protocone position relative to the hypocone: (0) lin- cone: (0) anterior; (1) posterior (Matthew, 1929; Rich, gual; (1) labial; (2) aligned (Storch and Qiu, 1991). 1981; Butler, 1948). 80. MI, protecone base: (0) anteromedially expanded to-
58. P4, carnassiform notch: (0) present; (1) absent (Rich, ward the P4; (1) not (Stevens, 1977).
1981). 81. MI, paracone shape: (0) conical; (1) crescentic (Munthe
59. P4, lingual roots: (0) one; (1) two fused; (2) two (But- and West, 1980). ler, 1948; Frost et al., 1991; Storch and Qiu, 1991). 82. MI, paracone size relative to the metacone: (0) approx-
60. P4, hypocone position relative to the protocone: (0) lin- imately equal; (1) smaller; (2) larger (Munthe and West, gual; (1) labial; (2) aligned (Storch and Qiu, 1991), 1980).
61. P4, link between protocone and hypocone: (0) crest; (1) 83. M1, paracone height: (0) largest cusp; (1) second largest adjoined by base proximity, no crest; (2) hypocone iso- cusp; (2) third largest cusp; (3) smallest cusp; (4) all lated (Matthew, 1929; Rich, 1981). cusps approximately equal in size (Matthews, 1929.
62. P4, metastyle: (0) high; (1) low; (2) absent (Stevens, Stevens, 1977; Rich, 1981).
1977; Butler, 1948). 84. M1, paracone position relative to the metacone: (0) labi-
63. Pd, parastyle: (0) distinct; (1) weak; (2) absent (Matthew, al; (1) lingual; (2) aligned (Koerner, 1940).
1929; Butler, 1948; Stevens, 1977; Black et al., 1980; 85. M1, centrocrista: (0) present; (1) absent (Rich, 1981). Rich, 1981). 86. M1, preparacrista: (0) strong; (1) weak; (2) absent (Rich,
64, P4, anterior cingulum: (0) present; (1) absent; (2) par- 1981). tial (Matthew, 1929; Rich, 1981). 87. M1, metacone shape: (0) crescentic; (1) conical (Munthe
65. P4, cingulum: (0) extends around hypocone; (1) not; (2) and West, 1980). absent (Butler, 1948). 88. M1, hypocone height relative to all of the other cusps:
66. P4, labial cingulum: (0) present; (1) absent (Matthew, (Q) tallest; (1) shortest; (2) approximately equal (Rich, 1929; Rich, 1981). 1981; Butler, 1948; Storch and Qiu, 1991).
67. P4/M1 position: (0) oblique to tooth row; (1) not (But- 89. M1, paraconule: (0) present; (1) absent (Matthews, ler, 1948). 1929; Munthe and West, 1980; Rich, 1981; Butler,
68. M1, size: (0) largest tooth of dentary; (1) not (Matthew, 1948).
1929; Munthe and West, 1980; Rich, 1981; Storch and 90. M1, preparaconule crista: (0) well developed; (1) not Qiu, 1991). (Rich, 1981).
69. M1, shape: (0) transversely rectangle; (1) antero-poste- 91. M1, preprotocrista: (0) well developed; (1) not (Rich, riorly rectangle; (2) quadrate (de Blainville, 1840; Koer- 1981). ner, 1940; Butler, 1948, 1988; Crusafonte et al., 1955; 92. M1, crest between protocone and metaconule: (0) pres- Stevens, 1977; Munthe and West, 1980; Rich, 1981; ent; (1) absent (Rich, 1981; Butler, 1948; Storch and Storch and Qiu, 1991). Qiu, 1991).
70. M1, anterior border: (0) straight; (1) concave; (2) con- 93. M1, hypocone: (0) crest joins it to the protocone-meta- vex (Rich, 1981). conule crest; (1) not (isolated) (Matthews, 1929; Munthe
71. M1, lingual roots: (0) separate; (1) fused (Storch and and West, 1980; Rich, 1981; Butler, 1948; Storch and Qiu, 1991). Qiu, 1991).
72. M1, metaconule: (0) present; (1) absent (Matthew, 1929, 94. M1, metastyle: (0) present; (1) absent (Viret, 1938; Viret, 1938; Butler, 1948, 1988; Black et al., 1980; Storch and Qiu, 1991).
Munthe and West, 1980; Storch and Qiu, 1991). 95. M1, metastyle apex: (0) high; (1) low (Munthe and West,
73. M1, metaconule: (Q) isolated; (1) not (Matthew, 1929, 1980).
Viret, 1938; Butler, 1948, 1988; Black et al., 1980; 96. M1, metastyle position relative to the metacone: (0) labi- Munthe and West, 1980; Storch and Qiu, 1991), al; (1) posterior (Butler, 1948).
74. M1, metaconule shape: (0) conical; (1) elliptical; (2) 97. M1, mesostyle: (0) present; (1) absent (Munthe and crescentic (Munthe and West, 1980). West, 1980; Rich, 1981).
75. MI, postmetaconule crista extension: (0) to the meta- 98. M1, parastyle: (0) present; (1) absent (Black et al., cone; (1) not; (2) absent (Matthew, 1929; Rich, 1981). 1980).
76. M1, protocone height: (0) tallest cusp; (1) second tallest 99. M1, metacrista: (0) present; (1) absent (Matthews, 1929; cusp; (2) third tallest cusp; (3) approximately equal in Munthe and West, 1980; Rich, 1981). height to all other cusps (Stevens, 1977; Rich, 1981). 160. M1, cingulum: (0) surrounds tooth; (1) discontinuous;
77. M1, protocone shape: (0) crescentic; (1) conical (2) absent (Matthews, 1929; Rich, 1981).
(Munthe and West, 1980). 101. M1, lingual cingulum: (0) present; (1) absent; (2) bead-
78. M1, protocone position relative to the paracone: (0) ante- ing (Stevens, 1977; Munthe and West, 1980; Rich, 1981). rior; (1) posterior; (2) equivalent (Matthews, 1929; 102. M1, anterior cingulum: (0) present; (1) absent (Stevens,
Stevens, 1977; Rich, 1981).
1977, Munthe and West, 1980; Rich, 1981).
29
30 AMERICAN MUSEUM NOVITATES NO. 3340
APPENDIX 1 Continued
103.
130.
M1, postcingulum: (0) present; (1) absent (Stevens,
104.
105.
106.
107.
108.
109,
110.
111.
112,
113.
114.
115.
116.
117.
118.
119.
120.
121.
122.
123.
124,
125.
126.
127,
128. 129.
1977; Munthe and West, 1980; Rich, 1981).
M1, labial cingulum: (0) present; (1) absent (Matthew, 1929; Stevens, 1977; Rich, 1981).
M2, shape: (0) transversely rectangle; (1) antero-poste- riorly rectangle; (2) quadrate (Koemer, 1940; Black et al., 1980; Storch and Qiu, 1991).
M2, lingual roots: (0) fused; (1) separate (Butler, 1948; Black et al., 1980; Frost et al., 1991; Gould, 1995). M2, anterior margins: (0) convex; (1) concave; (2) straight (Black et al., 1980).
M2, posterior margin: (0) convex; (1) concave; (2) straight (Black et al., 1980).
M2, protocone size relative to the paracone: (0) equal; (1) larger; (2) smaller.
M2, paracone position relative to the metacone: (0) lin- gual; (1) labial; (2) aligned (Black et al., 1980).
M2, hypocone: (0) isolated; (1) not (Matthews, 1929; Munthe and West, 1980; Rich, 1981; Butler, 1948; Storch and Qiu, 1991).
M2, metaconule: (0) present; (1) absent (Black et al., 1980; Munthe and West, 1980).
M2, metaconule: (0) isolated; (1) not (Black et al., 1980; Storch and Qiu, 1991).
M2, paraconule: (0) present; (1) absent (Black et al., 1980; Munthe and West, 1980).
M2, metaconule postion relative to the paraconule: (0) labial; (1) lingual (Black et al., 1980).
M2, metaconule size: (0) twice the size of the para- conule; (1) not (Black et al., 1980).
M2, metastyle: (0) present; (1) absent (Black et al., 1980; Munthe and West, 1980).
M2, parastyle: (0) present; (1) absent (Black et al., 1980, Munthe and West, 1980).
M2, mesostyle: (0) present; (1) absent (Black et ai., 1980).
M2, posthypocrista: (0) present; (1) absent (Black et al., 1980; Storch and Qiu, 1991).
M2, posthypocrista extension: (0) to postctngulum, (1) not (Black et al., 1980).
M2, preprotocrista extension: (0) to paraconule; (1) not (Black et al., 1986).
M2, preprotocrista extension: (0) to paracone; (1) not (Matthews, 1929; Rich, 1981).
M2, lingual cingulum: (0) present; (1) absent (Rich, 1981).
M2, anterior cingulum: (0) distinct; (1) partial; (2) absent (Rich, 1981).
M2, labial cingulum: (0) present; (1) absent; (2) partial (Rich, 1981).
M2, posterior cingulum: (0) present, (1) absent, (2) par- tial (Rich, 1981).
M2, cingula condition: (0) weak; (1) strong (Rich, 1981). M3: (0) present; (1) absent (Rich, 1981; Novacek, 1985, Novacek et al., 1985; Gould, 1995).
131.
132.
133. 134.
135.
136.
137.
138.
139.
140. 141.
142.
143.
144.
145.
146.
147.
148.
149,
150.
151.
152.
153.
154.
M3, roots: (0) four; (1) three; (2) two separate; (3) two fused (Butler, 1948; Rich, 1981; Frost et al., 1991; Gould, 1995).
M3, hypocone: (0) present, sits on cingulum; (1) absent; (2) fused to metacone (= metastylar spur) (Koerner, 1940; Butler, 1948: Munthe and West, 1980; Novacek, 1985; Novacek et al., 1985; Frost et al., 1991; Storch and Qiu, 1991; Gould, 1995).
M3, metacone: (0) large; (1) small; (2) absent (Koerner, 1940; Butler, 1948; Munthe and West, 1980; Novacek, 1985; Novacek et al., 1985; Frost et al., 1991; Storch and Qiu, 1991; Gould, 1995),
M3, protocone size; (0) large; (1) small.
M3, main cusps: (0) equally developed; (1) not (Munthe and West, 1980),
M3, metaconule: (0) present; (1) absent (Butler, 1948; Munthe and West, 1980).
M3, parastyle: (0) present; (1) absent (Munthe and West, 1980).
M3, anterior cingulum: (0) present; (1) absent (Munthe and West, 1980).
M3, posterolingual cingulum: (0) present; (1) absent (Munthe and West, 1980).
M3, posterolabial cingulum: (0) present; (1) absent, (2) partial (Rich, 1981).
M3, posterior cingulum: (0) present; (1) absent.
il: (0) present; (1) absent (Leche, 1902; Butler, 1948, 1988; Stevens, 1977; Rich, 1981; Frost et al., 1991). il, size relative to i2: (0) approximately equal; (1) larg- er (Butler, 1948; Novacek, 1986; Rich, 1981; Frost et al., 1991; Storch and Qiu, 1991).
il, shape: (0) spatulate; (1) conical (Rich, 1981).
il, root: (G) short; (1) long (Rich and Rasmussen, 1973).
12: (O) present; (1) absent (Rich, 1981).
i2, size: (O) enlarged; (1) reduced (Butler, 1948, 1988; Novacek, 1985; Novacek et al., 1985; Frost et al., 1991; Gould, 1995).
i2, shape: (0) spatulate; (1) conical (Stevens, 1977; Munthe and West, 1980).
i2, position: (0) overlaps preceding tooth; (1) not (Rich, 1981).
12, anterior midline crest: (0) ends posterior to proto- conid; (1) not (Rich, 1981),
13: (0) present; (1) absent (Butler, 1948).
13, size relative to other incisors: (0) smaller; (1) approx- imately equal; (2) larger (Butler, 1948, 1988; Corbet, 1988; Frost et al., 1991).
Lower canine, size relative to pl: (0) approximately equal; (1) greater (Butler, 1948; Rich, 1981; Frost et al., 1991; Storch and Qiu, 1991).
Lower canine mophology: (0) like 12/p2; (1) not (Storch and Qiu, 1991).
Lower canine: (0) overlaps preceding tooth; (1) not (Rich, 1981).
2001
155. 156. 157. 158. 159, 160. 161, 162.
163.
164.
165. 166. 167. 168. 169. 170. 171. 172. 173. 174. 175,
176.
177. 178.
179.
180.
181.
182.
183.
GOULD: DENTAL MORPHOLOGY OF HEDGEHOGS
APPENDIX 1 Continued
Lower canine, anterior midline crest: (0) present; (1) 184. p4, paraconid: (QO) strong; (1) weak; (2) absent (Matthew, absent (Rich, 1981). 1929; Rich, 1981; Storch and Qiu, 1991). Lower canine, anterior midline crest: (0) ends posterior 185. p4, paraconid height relative to protoconid: (0) approxi- to principal cusp; (1) not (Rich, 1981). mately equal; (1) shorter (Koerner, 1940; Butler, 1948, Lower canine, posterior ridge: (0) present; (1) absent; 1988; Black et al., 1980; Munthe and West, 1980; Rich, (2) weak (Rich, 1981). 1981). Lower canine, lingual ridge: (0) present; (1) absent (Rich, 186. p4, paraconid position relative to protoconid: (0) an- 1981). terolingual; (1) directly anterior (Stevens, 1977). Lower canine, basal cuspule: (0) present; (1) absent (Frost 187. p4, paraconid: (0) separated from protoconid by notch; et al., 1991; Gould, 1995). (1) not (Stevens, 1977). pl: (0) present; (1) absent (Butler, 1948, 1988, Rich, 188. p4, protoconid position: (0) cental; (1) labial (Munthe 1981; Frost et al., 1991; Storch and Qiu, 1991). and West, 1980). pl, roots: (0) single; (1) partly divided (Butler, 1948). 189. p4, protoconid size: (0) greater than metaconid; (1) not pl, cuspules: (0) one; (1) two; (2) three. (Rich, 1981; Butler, 1948). p2: (0) present; (1) absent (Rich, 1981; Novacek, 1985; 190. p4, metaconid: (0) present; (1) absent (Matthew, 1929; Novacek et al., 1985; Gould, 1995). Rich, 1981). p2, roots: (0) one; (1) two (Koerner, 1940; Butler, 1948, 191. pd, metaconid size: (0) small; (1) large (Butler, 1948, 1988: Black et al., 1980; Munthe and West, 1980; Rich, 1988; Stevens, 1977; Munthe and West, 1980; Storch and 1981). Qiu, 1991). p2, cuspules: (0) one; (1) two; (2) three; (3) four; (4) 192. p4, posterior talonid cuspule(s): (0) present; (1) absent absent. (Stevens, 1977; Rich, 1981: Butler, 1948). p2, cingulum: (0) present; (1) absent; (2) partial (Munthe 193. p4, number of posterior talonid cuspules: (0) one; (1) and West, 1980). two; (2) three (Stevens, 1977; Rich, 1981; Butler, 1948). p2, position: (0) overlaps preceding tooth; (1) not (Rich, 194, p4, cingulum: (0) strong; (1) weak; (2) absent (Rich, 1981). 1981). p2, anterior midline crest: (0) present; (1) absent (Rich, 195. p4, size relative to m1: (0) approximately equal: (1) 1981). smaller (Storch and Qiu, 1991). p2, anterior midline crest: (0) ends posterior to proto- 196. Prevallid shear: (0) present; (1) absent (Stevens, 1977; conid; (1) not (Rich, 1981). Novacek, 1985, 1986). p2, lingual ridge: (0) present; (1) absent (Rich, 1981). 197. ml, postcingulum: (0) strong; (1) weak; (2) absent (Black p2, posterior ridge: (0) present; (1) absent (Rich, 1981). et al., 1980; Rich, 1981; Storch and Qiu, 1991). p3: (Q) present; (1) absent (Butler, 1948; Rich, 1981). 198. ml, trigonids: (0) high, short talonid; (1) low, talonid p3, roots: (0) two; (1) one; (2) two fused (Butler, 1948; expanded (Matthew, 1929; Stevens, 1977; Rich, 1981; Novacek, 1985; Novacek et al., 1985; Corbet, 1988; Butler, {948; Novacek, 1985; Novacek et al., 1985; Frost Storch and Qiu, 1991). et al., 1991; Gould, 1995). p3, size relative to p2: (0) much larger; (1) approximate- 199. ml, protoconid: (0) lingually inclined; (1) not (Black et ly equal (Butler, 1948; Munthe and West, 1980). al., 1980). p3, cusps: (0) two; (1) one; (2) three (Munthe and West, 200. ml, protocristid: (0) contacts metaconid and protoconid; 1980). (1) no contact; (2) absent. p3, posterior margin; (0) wide; (1) narrow (Munthe and 201. ml, metaconid position: (0) anterior to protoconid; (1) not West, 1980). (Matthew, 1929; Rich, 1981; Butler, 1948). p3, metaconid crest: (0) present; (1) absent (Butler, 1948). 202. ml, metaconid height relative to paraconid: (0) greater; p3, posterolingual cusp: (0) prominent; (1) weak or (1) approximately equal or smaller (Black et al., 1980). absent (Munthe and West, 1980). 203. ml, paraconid: (0) large; (1) small; (2) absent (Rich and p3, cingulum: (0) present; (1) absent; (2) partial (Munthe Rasmussen, 1973; Rich, 1981). and West, 1980). 204. ml, labial wall: (0) markedly concave; (1) not (Matthew, p4, talonid: (0) elongated; (1) short (Novacek, 1985; 1929; Rich, 1981). Novacek et al., 1985; Gould, 1995). 205. m1, lingual wall: (0) markedly concave; (1) not (Mat- p4, talonid: (0) greatest breadth of tooth; (1) not (Matthew, thew, 1929; Rich, 1981). 1929; Stevens, 1977; Rich, 1981; Butler, 1948). 206. ml, talonid: (0) enclosed lingually by entocristid; (1) not p4, talonid posterior ridge: (0) present; (1) absent (Mat- (Rich, 1981; Storch and Qiu, 1991). thew, 1929; Rich, 1981; Storch and Qiu, 1991}, 207. ml, talonid: (0) opens posteriorly; (1) closed (Stevens, p4, posterolabial cusp: (0) present; (1) absent (Butler, 1977). 1948). 208. ml, hypoconid: (0) isolated; (1) not (Black et al., 1980).
31
52
209.
210.
211.
212, 213.
214.
215,
216.
217,
218.
219.
220.
221,
ded,
223:
224.
225.
226. a On
228.
AMERICAN MUSEUM NOVITATES
NO. 3340
APPENDIX 1 Continued
ml, entoconid size: (0) larger than the hypoconid; (1) 229, larger than the paraconid; (2) approximately equal to all other cusps; (3) larger than the hypoconid and paraconid 230. (Matthew, 1929; Butler, 1948; Stevens, 1977; Rich, 1981). 231. ml, entostylid: (0) present; (1) absent (Black et al., 1980). peas m1, entocristid: (0) high; (1) low; (2) absent (Rich, 1981; 233. Butler, 1948; Storch and Quu, 1991). m1, hypoconulid: (0) present; (1) absent (Butler, 1948). 234. ml, cristid obliqua orientation: (0) antero-posteriorly directed; (1) inclined; (2) absent (Black et al., 1980). 235. ml, hypocristid: (0) extends to posterior cingulum; (1) not; (2) absent (Engesser, 1972; Black et al., 1980). 236. ml, labial cingulum: (0) continuous around hypoconid; (1) not (Stevens, 1977; Rich, 1981; Butler, 1948), ml, labial cingulum: (0) strong; (1) weak; (2) absent (Matthew, 1929; Rich, 1981; Novacek, 1985; Novacek 237. et al., 1985; Gould, 1995). ml, entoconulid: (0) present; (1) absent (Black et al., 238. 1980). 239. ml, paraconid position relative to the metaconid: (0) lingual; (1) labial; (2) aligned. 240. ml, paraconid position relative to the protoconid: (0) lingual; (1) labial; (2) aligned. ml, paracristid: (0) horizontal; (1) attenuated. 241. m1, paracristid lingual extension: (0) terminates at mid- anterior of tooth; (1) extends to lingual border (Rich, 242. 1981; Butler, 1948). m2, size relative to m1: (0) smaller; (1) larger (Matthew, 243. 1929: Stevens, 1977; Rich, 1981; Butler, 1948). m2, trigonids: (0) high, short talonid; (1) low, expand- ed talonid (Stevens, 1977; Rich, 1981). 244. m2, protoconid height relative to the metaconid: (0) smaller; (1) approximately equal; (2) larger (Matthew, 245. 1929: Rich, 1981). 246. m2, paraconid: (0) present; (1) absent (Rich, 1981; Storch and Qiu, 1991). m2, paraconid shelf: (0) present; (1) absent. 247. m2, paraconid swelling: (0) present; (1) absent (Rich, 1981). m2, paracristid lingual extension: (0) approximately 248. equal to m1; (1) not; (2) more lingual than m1 (Butler, 249.
1948).
m2, entostylid: (0) absent; (1) weak; (2) strong (Black et al., 1980).
m2, posterior cingulum: (0) present; (1) absent (Rich, 1981).
m2, posterior cingulum: (0) connects to entoconid; (1) not (Rich, 1981; Storch and Qiu, 1991).
m2, entoconulid: (0) present; (1) absent.
m2, paraconid position relative to the metaconid: (0) lingual; (1) labial; (2) aligned.
m2, paraconid position relative to the protoconid: (0) lingual; (1) labial; (2) aligned.
m2, hypoconid position relative to the protoconid: (0) lingual; (1) labial; (2) aligned (Rich, 1981).
m2, entoconid size: (0) approximately equal to the pro- toconid; (1) tallest cusp; (2) approximately equal to the hypoconid; (3) approximately equal to the metaconid (Matthew, 1929, Rich, 1981).
m2, talonid: (0) posteriorly narrow; (1) not (Matthew, 1929; Rich, 1981).
m2, talonid basin: (0) lingually enclosed; (1) not.
m2, hypoconulid: (0) present; (1) absent (Rich, 1981; Black et al., 1980).
m2, cristid obliqua contact point: (0) at the base of the protoconid; (1) midheight of protoconid (Matthew, 1929; Rich, 1981).
m2, entocristid: (0) high; (1) low; (2) absent (Rich, 1981; Storch and Qiu, 1991).
m2, labial cingulum: (0) continuous; (1) discontinuous; (2) absent (Matthew, 1929; Rich, 1981),
m3: (0) present; (1) absent (Rich and Rasmussen, 1973; Stevens, 1977; Rich, 1981; Novacek, 1985; Novacek et al., 1985; Gould, 1995).
m3, paraconid: (0) swollen; (1) normal; (2) absent (Rich, 1981; Storch and Qiu, 1991).
m3, paraconid shelf: (0) present; (1) absent.
m3, postcingulum: (0) present; (1) absent (Koemer, 1940; Rich and Rasmussen, 1973; Rich, 1981; Storch and Qiu, 1991),
m3, talonid: (0) present; (1) absent (Koerner, 1940; Rich and Rasmussen, 1973; Munthe and West, 1980; Rich, 1981; Butler, 1948; Frost et al., 1991; Gould, 1995). m3, trigonids: (0) high; (1) short.
m3, lingual cingulum: (0) present; (1) absent (Munthe and West, 1980).
2001
Atelerix
Atelerix albiventris (25)
USNM USNM USNM USNM USNM USNM USNM USNM USNM USNM USNM USNM USNM USNM USNM USNM USNM USNM USNM USNM USNM USNM USNM USNM USNM
378723 378725 378726 378728 378729 378730 378731 378732 378740 378741 378742 378746 378747 378748 378750 378751 378752 402179 402180 402181 402182 402183 402184 375927 375928
N7eszeZztrtretitreszeZS3 Td TAINS
Atelerix algirus (21)
USNM USNM USNM USNM USNM USNM USNM USNM USNM USNM USNM USNM USNM USNM USNM USNM USNM USNM USNM USNM USNM
476050 476051 476052 476053 476054 476055 476056 476057 476058 476059 476060 476061 476062 476063 476064 476065 476066 470578 470579 482681 140766
~Zquecwzttzsettrezenyrnnes Day
GOULD: DENTAL MORPHOLOGY OF HEDGEHOGS 33
APPENDIX 2
Specimens Reviewed in Discrete Dental Analyses Abbreviations: AMNH = American Museum of Natural History; USNM = United States National Museum (Smithsonian); F = female; M = male; (#) = number of specimens reviewed.
Nigeria, Kano Prov. Nigeria, Kano Prov. Nigeria, NW Zaria Nigeria, NW Zaria Nigeria, NW Zaria Nigeria, NW Zaria Nigeria, N Sokoto Nigeria, N Sokoto Nigeria, N Sokoto Nigeria, N Sokoto Nigeria, N Sokoto Nigeria, N Sokoto Nigeria, N Sokoto Nigeria, N Sokoto Nigeria, not labeled Nigeria, N Sokoto Nigeria, N Sokoto Nigeria, N Sokoto Nigeria, N Sokoto Nigeria, N Sokoto Nigeria, N Sokoto Nigeria, N Sokoto Nigeria, N Sokoto Nigeria, Plateau Prov. Nigeria, S Kabwir
Morocco, Fes Prov. Morocco, Oujda Prov. Morocco, Ksar Es Souk Prov. Morocco, Ksar Es Souk Prov. Morocco, Fes Prov. Morocco, Fes Prov. Morocco, Al Hoceima Prov. Morocco, Oujda Prov. Morocco, Agadir Prov. Morocco, Adadir Prov. Morocco, Adadir Prov. Morocco, Adadir Prov. Morocco, Tetouan Prov, Morocco, Oujda Prov. Morocco, Ksar Es Souk Prov. Morocco, Ksar Es Souk Prov. Morocco, Beni-Mellal Prov. Morocco, Agadir Prov. Morocco, Agadir Prov. Morocco, Agadir Prov.
Sud Tunis, Djerba
Echinosorex (32)
Echinosorex gymnurus
USNM USNM USNM USNM USNM USNM USNM USNM USNM USNM USNM USNM USNM USNM USNM USNM USNM USNM USNM USNM USNM USNM USNM USNM USNM USNM
487885 487887 487888 487889 487890 487901 487892 487893 487894 487895 487896 487897 487898 487899 487900 487902 487903 283474 283475 115489 357885 3787
357886 367888 487886 357887
SUEZ VE VIIBVME NEES ECVE NUE 7
west Malaysia west Malaysia west Malaysia west Malaysia west Malaysia west Malaysia west Malaysia west Malaysia west Malaysia west Malaysia west Malaysia west Malaysia west Malaysia west Malaysia west Malaysia west Malaysia west Malaysia Malaya Malaya Pahang: Rumpin River Malaysia: Selangor Singapore Malaysia Malaysia Malaysia Malaysia
Echinosorex g. “dealbatus”
USNM
Echinosorex 2. “alba”
USNM USNM USNM USNM USNM
83704
145581 145582 145584 145585 145586
Erinaceus
MJ
Te Hn Ty
“Butaw” or Tikus Island, Sumatra
West Borneo, Sempang River West Borneo, Sempang River West Borneo, Sempang River West Borneo, Sempang River West Borneo, Sempang River
Erinaceus amurensis (11)
USNM USNM USNM USNM USNM USNM USNM USNM USNM USNM USNM
176251 199681 239770 239590 239591 239592 197779 270541 270542 240325 252158
M M
11s DN nNy-~
N. China, Tiensin
N. China, Tiensin
China, Ningpo
China, Hunan, Yochow
China, Hunan, Yochow
China, Hunan, Yochow
China, Kirin Prov.
Inner Mongolia, Grter. Khingan Inner Mongolia, Grter. Khingan China, Shanghai
China, Shanghai
34
AMERICAN MUSEUM NOVITATES
APPENDIX 2 Continued
Erinaceus europaeus (24) Hemiechinus aethiopicus (= Paraechinus, 25) (continued)
USNM USNM USNM USNM USNM USNM
USNM USNM USNM USNM USNM USNM USNM USNM USNM
USNM USNM USNM
USNM USNM USNM USNM USNM
USNM
153409 153410 153411 153412 1856
186556
251763 251764 251765 251766 251767 251768 271142 151668 37465 (12244) 85619 86923 36034 (20807) 174660 794 795 34959 (19246) 34960 (19247) 84739
Hemiechinus
ie ee
NZ MEEZEUUMY
ig Beg
Ts
F
Wales, Cardiff
England, Wandsworths Comn, Bavaria, Strass
Germany, Ingelheim
Bavaria
W. Germany, Braunschweig (Saxony)
Germany
Germany
Germany
Germany
Germany
Germany
Germany
W. Germany, Baden Wurtemburg Germany, Heidelburg
Germany, Braunschweig Ireland, Glenmore County England
Channel Islands, Guermsey England England England (Nat. Zool. Park) England (Nat. Zool. Park)
Switzerland, St. Gallen
Hemiechinus aethiopicus (= Paraechinus, 25)
USNM USNM USNM USNM USNM USNM
USNM USNM USNM USNM USNM USNM USNM USNM USNM USNM USNM USNM USNM USNM
311732 311737 311738 311739 311740 321572
325906 325907 325908 384832 410872 410873 482512 470563 470564 470565 470566 470567 470568 470569
cag: i
MZZZUWUVEZETEEAET
Egypt, Sudan Admin. Area Egypt, Western Desert, Gov. Egypt, Western Desert, Gov. Egypt, Sinai, St. Catherine’s Egypt, Sinai, St. Catherine’s SW Saudi Arabia, E. Aden Protectorate, Taribin
Egypt, Sinai
Egypt, St. Catherine’s Egypt, St. Catherine’s Mauritania, Atar Mauritania, Kiffa Mauritania, Kiffa
Niger, 5 km NE Agadez Morocco, Agadir Prov. Morocco, Agadir Prov. Morocco, Tarfaya Prov. Morocco, Ouarzazate Prov. Morocco, Quarzazate Prov. Morocco, Quarzazate Prov. Morocco, Ouarzazate Prov.
USNM USNM USNM USNM USNM
NO. 3340
476067 476068 476069 482862 482863
F
M M M
Morocco, Ksar Es Souk Prov. Morocco, Ksar Es Souk Prov. Morocco, Ksar Es Souk Prov. Morocco, Agadir Prov. Morocco, Agadir Prov.
Hemiechinus auritus (25)
AMNH AMNH AMNH AMNH AMNH AMNH AMNH AMNH AMNH AMNH AMNH AMNH AMNH AMNH AMNH AMNH AMNH AMNH AMNH AMNH AMNH
AMNH AMNH AMNH AMNH
203197 203198 203199 203200 170226 170227 170228 170229 244379 244380 244384 176282 87085 85309 85308 31248 57216 57217 57222 84001 31246
184065 22876 22889 80021
x
NMer~n~ Zerg ut
2255 a 12S
? ? i ?
Egypt, Giza, Imbaba, Kafr Hakem Egypt, Giza, Imbaba, Manshiyet Egypt, Giza, Imbaba, Manshiyet Egypt, Giza, Imbaba, Tanash Pakistan, Baluchistan, Quetta Pakistan, Baluchistan, Quetta Pakistan, Baluchistan, Quetta Pakistan, Baluchistan, Quetta Pakistan, Baluchistan, Kalat Pakistan, Baluchistan, Kalat Pakistan, Baluchistan, Quetta USSR Turkmenskaya SSR USSR, Uzbekskaya SSR Fergana USSR, Kazakhsakay SSR USSR, Kazakhsakay SSR USSR, RS FSR Sarepta, NYZS Mongolia, Oyor-Hangay Prov. Mongolia, Oyor-Hangay Prov. Mongolia, Oyor-Hangay Prov. Mongolia, Oyor-Hangay Prov. China, Xinjiang Uygur Zizhiqu Kashi
Israel, Kvutzat-urim Zoo NYSZS
NYSZS
NYSZS
Hemiechinus hypomelas (19)
USNM USNM USNM USNM USNM USNM USNM USNM USNM USNM USNM USNM USNM USNM USNM USNM USNM USNM USNM
326695 326696 326697 326698 326699 327914 326700 326701 327913 327915 4529511 368931 368932 368933 368934 368935 368936 368937 410929
|
ae ee 7 Ue ee Se eee Ss
Iran, Khursan, Turbat-]-Haibari Iran, Khursan, Turbat-I-Haiban Iran, Khursan, Turbat-I-Haibari Iran, Khursan, Turbat-I-Haibari Iran, Khursan, Turbat-I-Haiban Iran, Dasnt-I-Lut Majak
Iran, 6 km N of Kashmar
Iran, Turbat-I-Haibari
Iran, Dasnt-]-Lut Majak
Iran, Majan
Pakistan, Gulistan Quetta Dist. Pakistan, Muzaffargarh Alipur Pakistan, Deragazikahn Pakistan, Deragazikahn Pakistan, Muzaffargarh Alipur Pakistan, Muzaffargarh Alipur Pakistan, Muzaffargarh Alipur Pakistan, Muzaffargarh Alipur Pakistan, Deragazikahn
2001 GOULD: DENTAL MORPHOLOGY OF HEDGEHOGS
APPENDIX 2 Continued
Hylomys Hylomys sinenesis (25) (continued) Hylomys sinenesis (25) AMNH 44268 M_— China, Taiping-pu, Yun-nan Prov. AMNH 115505. F N. Burma, Kachin Prov. AMNH 44270 F China, Taiping-pu, Yun-nan Prov. AMNH 115506 M_N. Burma, Kachin Prov. AMNH 44271 M_— China, Taiping-pu, Yun-nan Prov. AMNH 115508 F N. Burma, Kachin Prov. AMNH 57199? China, Yun-nan Prov. AMNH 115509 F N., Burma, Kachin Prov. AMNH 115510 M_ N. Burma, Kachin Prov. Hylomys suillus (16) AMNH 115511 F N. Burma, Kachin Prov. USNM 481278 F Java AMNH 115512 M __ N. Burma, Kachin Prov. USNM 481279 F Java AMNH 115514 M_ N. Burma, Kachin Prov. USNM 481280 F — Java AMNH 115515 F N. Burma, Kachin Prov, USNM 481281 F Java AMNH 115516 F _ N. Burma, Kachin Prov. USNM 481283 F Java AMNH 115517 M _ N. Burma, Kachin Prov. USNM 481284 F Java AMNH 115518 M N. Burma, Kachin Prov. USNM 481285 F Java AMNH 115519 M_ N. Burma, Kachin Prov. USNM 481286 F — Java AMNH 115520 M _N. Burma, Kachin Prov. USNM 481287 M — Java AMNH 115522 F N. Burma, Kachin Prov. USNM 481288 M — Java AMNH 115523 M_ N. Burma, Kachin Prov. USNM 481289 F — Java AMNH 115524 MN, Burma, Kachin Prov. USNM 481290 F Java AMNH 115525 M _ N. Burma, Kachin Prov. USNM 521659 M | Java AMNH 44248 F — China, Mu-cheng, Yun-nan Prov. USNM 521660 M | Java AMNH 44249 M_ China, Mu-cheng, Yun-nan Prov. USNM 521661 M — Java AMNH 44267 M_— China, Taiping-pu, Yun-nan Prov. USNM 155660 F — Java
36 AMERICAN MUSEUM NOVITATES NO. 3340
APPENDIX 3
Frequency Distribution of 246 Transformation Series across 10 Taxa Abbreviations: N = sample size; M = male; F = female; ? = sex undetermined; J = juvenile; MA = mature adult; W = worn; TS# = transformation series number (see appendix 1); column headings 0, 1, 2, 3, 4 = character state numbers (see appendix 1); A = assymetrical character expression; T = total number of specimens scored.
Echinosorex gymnura Hylomys sinensis Hylomys suillus Atelerix albiventris Atelerix algirus (N = 32; M-14, F-16, 9-2; | (N = 25; M-15, F-9, ?-1; (N = 16; M-5, F-11; (N = 25; M-11, F-14; (N = 21; M-7, F-13, 7-1; J-7, MA-18, W-7) J-0, MA-24, W-1) J-3, MA-11, W-2) J-4, MA-13, W-8) J-5, MA-7, W-9)
TS#! 0 12 3 4 A Tj 0 12 3 4 AT} O0O 1 2 3 4 AT] O 123 4 A O 123 4 AT 4)27 5 32 | 25 25 | 16 16 | 25 21 21 5) 32 32 } 25 25| 6 10 16 25 2] 21 6 | 32 32 | 25 25 | 16 16 | 25 21 2] 7) 2 5 10 17 23 23} 2 2 10 2 16 25 25 20 1 21 8 | 27 27 | 25 25|12 4 16 25 25 21 2] 9 | 25 25 25 25 16 16 | 25 25 | 2] 21 10/30 1 31725 25116 16} 25 25 | 21 21 11 | 29 29 | 25 25 | 16 16 | 25 25; 114 #1 2 18 12 NA 25 25 NA | 24 24 | 20 20 13 | 26 26 24 24 16 16 25 25 21 21 14; 8 12 1 21 23 23 16 16] 12 6 18 15 15 15! 612 1 1 20/12 8 20; 3 13 16] 6 12 18! 6 9 15 16} 19 1 20 22 «1 23 16 16 25 25) 2 19 21 17 | 27 27 23 23 12. 2: 2 §6119 19 5 8 13 18! 31 31 18 4 22 14 2 16 25 25 19 19 19/24 2 5 31 3.19 22 | 16 16 | 24 24 | 16 1 17 20 | 32 32/16 5 21 | 16 16! 5 16 1 22 19 19 21 | 29 29 24 24 16 16! 3 20 23) 415 19 22) 29 29 24 24 16 16] 2 22 24 19 19 23 7 14 3 24 4 20 24,10 2 4 16); 7 6 9 224; 1 11 7 19 24 | 29 2 31 25 25) 14 1 1 16 25 25 21 2] 25! 214 2 2 20 NA 13 2 15 NA NA 26) 117 2 20 NA; 6 8 115 NA NA 27 | 30 1 31 NA/ 14 1 15 NA NA 28 | 31 31 | 24 24; 16 16 | 25 25 | 21 21 29 214 16 24 24 3 10 3 16] It 6 17 1 25 20 20 30; 112 2 1 16 16 5 21/15 1 16/10 6 6 2 24; 2 4 13 19 31 31 31 20 3 23) 4 8 3 15]24 1 25 | 16 1 1 18 32 | 32 32 | 25 25 | 16 16|22 2 1 25/21 21 33 [ 3 29 32 25 25 16 16 23 23) 3 18 21 34} 27 5 32 25 25 16 16} 8 12 3 23/17 4 21 35 24 8 32 25 25 14 2 16 1 22 23; 2 4 15 21 361 6 26 32 25 25 16 16] 1 22 23; 317 20 37 | 12 17 3 32 NA 16 16] 2 20 221 6 13 19 38 | 15 16 31 NA NA! 1 1 2;| <3 6 39 | 32 32 25 25 16 16 23 23 21 21 40 | 32 32 25 25 16 16/12 9 1 22/11 5 16 4l NA NA | NA 4 4 8} 5 1 1 1 8 42 NA NA | NA 8 8 1 11 43 | 32 32 | 25 251 16 16/12 1 7 20 | 21 21 44} 3] 31 | 25 25/13 2 1 16) 15 6 2 23) 21 21 45) 2 29 31 25 25 16 16 21 2 23 21 21 46 1 I 25 25 14 14 NA NA 47; 415 7 5 31 25 25 16 16 25 25 21 21 48 2 30 32 25 25) 5 8 3 16 2 21 1 23 515 20 49; 32 32 25 25 | 16 16 22 «1 23/19 2 21 30; 16 12 3 3) 25 25 16 16; 7 15 1 23} 1 20 21 S51} 2 2 NA NA NA NA 52 | 32 32 25 25 16 16] 10 13 1 24/14 3 17 53|26 4 2 32 25 25 | 16 16] 5 13 3 21 2] 21 54 | 32 32 | 25 25 | 16 16|22 2 1 25] 21 21 55 | 32 32 | 25 25 | 16 16 | 22 22 | 21 21
56 | 32 32 | 25 25| 1 15 16,10 7 3 20] 21 21
2001
GOULD: DENTAL MORPHOLOGY OF HEDGEHOGS
APPENDIX 3 Continued
Echinosorex gymnura
(N = 32; M-14, F-46, 9-2;
0
58 59
60 | 12 61] 2 62
63 | 29 64] 3 65 | 14 66 | 19 67
68 | 32 69
70 | 15 71 | 30 72 | 29 73) 9 74 | 13 75 | 23 76| 6 77 | 30 78 | 23 79
B0 | 28 81 | 30 82} 1 83
84
85 | 13 86| 5 87
88
89 | 1 90] 5 91 | 25 92] 11 93] 6 94 | 30 95
96 | 30 97
98] 8 99 | 29 100| 4 101] 8 102 | 31 103 | 31 104 | 30 105
106
107| 7 108 | 1 109 |
110
111 | 16 112 | 30
1
3 32
i 15 32
2 26 16 11 31
26
14 15 27 16 12
29
29 20
27
32
10 30
1]
19 29
12
32
25 21
24
J-7, MA-18, W-7) 23 4AT
— ee KD
wa
Atelerix albiventris (N = 25; M-1], F-14;
Aylomys sinensis Hylomys suillus (N = 25; M-15, F-9, 7-1; (N = 16; M-S, F-11; J-0, MA-24, W-1) J-3, MA-11, W-2)
0123 4 A 6 123 4A TY] 0 32 15 1 16 ; 23 31 | 25 25)15 1 16 | 24 32. 25 25 iP 11| 7 28; 5 4 9 7 25/15 1 161 3 17 NA 16 16 | 11 32 | 25 25 16 16 | 22 32 25 25| 8 7 1 16} 2 31; 9 11 1 4 25] 1 15 16 31 22. °2. i 25 16 16] 5 31) 7 14 3 1 25) [ 15 16) | 31 25 25 16 16 32 | 25 25 | 16 16 | 25 32 25 25 12 16 32,10 12 3 25/13 3 16] 11 30; 25 25 NA 29 | 25 25 | 15 15| 9 23; 3 20 i 24; 5 4 6 16 20 | 24 24) 3 9 2 14} 2 28 | 22 ] i 24 t5 i5 14 25 25 144 15 30 | 25 25] 15 15} 15 27 | 25 25 | 15 15119 24; 2 31 10 2 25 16 16 30 25 25 16 16 30 | 25 25116 16118 27 25 25 16 16 | 1i 19 24 24 14 14/10 29 19 6 25 16 16 | 10 28 | 25 25; 4 11 15} 2 26 8 15 1 24) 4 6 3 2 15 27 24 1 25 15 15| 1 22 25 25 15 15 25/18 4 2 24| 1 «(14 15| 3 13) 1:19 20; I ee 25 | 24 24} 14 1 i5| 14 22; 8 10 6 24] 3 10 2 15] 6 17/13 1 i115} 3 9 3 15] 6 30 | 25 25 | 16 16 | 25 29 | 25 25 15 15 30 | 25 25 | 16 16 | 25 29 25 25 15 15 30| 6 17 2 25| 16 16 | 15 29 | 25 25 | 15 15 | 24 31 25 25 16 16 31 25 25 16 16 31|18 6 1 25! 2 12 2 16| 9 31)24 1 25 16 16 | 16 30 | 25 25; 15 1 16| 9 32 25 25 16 16 32 25 25 NA 32;13 3 4 4 24) 16 16 | 25 32} 8 2 15 25 16 16 30 24 25 15 15 30 5 8 12 25 14 2 16 29 24 24 16 16 30 | 25 25) 12 113) 5
1
] ! 17 13
24 20 24 25
Fe kA Nh WwW
24 25
Il 13
18 13
25
23
20 22 13
13
25 22 25 25
Atelerix algirus (N = 21; M-7, F-13, 9-1;
37
J-4, MA-13, W-8) J-5, MA-7, W-9) 2 tte A TiO tke 2) 3-4 wv OT 24 | 21 21 25 | 21 21 24 21 21 5 22} 21 21 12 NA 24 21 2] 22 25/210 8 20 251 7 13 20 25+ 8 Il 2 21 25{ 6 15 2] 25 2] 21 25 | 21 21 25 25 2) 2] 8 24/19 2 21 2 21 21 13 | 20 20 7 19 1 20 6:19 | 20 & 11 19 1 20 12 12 21 21 15} 21 21 2 2) } 21 21 24 1 20 21 25 21 21 19 | 2! 2] 5 22 | 2) 2] 13 | 21 21 23 | 1 14 2 17 17 | t4 2 16 12 19 21 21 19 1 20 21 13 21 21 13} 2 9 ia S$} 1] 2 14] 19 19 10 | 21 21 11} 14 1 15 25 | 21 21 25 21 21 25 | 21 21 24 21 21 23) 8 9 3 20 24 | 21 21 5 25 21 21 23) 6 5 9 1 21 23 | 20 20 23 | 21 2] 23! 21 21 25 25 21 21 NA | 2] 21 25 | 21 2) 25 21 2] 22 21 21 25 21 2] 25 21 2] 7/15 3 16
38
TS#
113 114 115 116 Hi? 118 119 120 121 122 123 124 125 126 127 128 129 130 131 132 133 134 135 136 137 138 139 140 141 142 143 144 145 146 147 148 149 150 151 152 153 154 155 156 157 158 159 160 161 162 163 164 165 166 167 168
Echinosorex gymnura (N = 32; M-14, F-16, ?-2;
AMERICAN MUSEUM NOVITATES
Hylomys sinensis (N = 25; M-15, F-9, 3-1; J-0, MA-24, W-1)
J-7, MA-18, W-7) 0123 44 6 22 1 9 19 1 10 6 4 31 16 i4 1 3] 1 28 11 27° 2 4 28 28 2 31 30 14 16 32 1 31 27 3 29 29 29 28 28 22 «6 28 24 4 28 30 30 2305 28 616 7 29 2 27 29 32 32 31 31 32 32 31 31 32 32 32 32 32 32 20 1 21 25 25 Az: 32 31 1 32 3] 31 30 30 32 32 4 27 31 5 5 422 4 30 30 30 10 19 1 30 29 2 1 32 301 31 It 5 1 1 1 19 30 2 32 29 29 19 3 2 } 25 4 4 15 1 24 26 26 23 2 25
25 25 25 25 25 25
4 20 1 25 20 4 24 7 18 25 25 25 25 25
2 23 25 2 2
2 2 2; 2
25 25 2302 25 25 25 8 10 18 1 24 25 235 25 24 1 25 NA
1 24 25 3 21 24 13. 9 1 23 13 «1 14 910 4 23 1 22 23 1 24 24 25 25 NA
NA
25 25 25 25 2 4 17 23 23 23 22 22 219 21
APPENDIX 3 Continued
Hylomys suillus
(N = 16; M-5, F-11;
J-3, MA-11, W-2) 012 3 4AT 1 9 3°13 13 13 NA NA 16 16 ll 4 15 16 16 5 1l 16 NA NA 16 16 16 16 4 9 2 15 14 2 16 16 16 12 2 14 16 16 16 16 16 16 14 2 16 16 16 16 16 15 15 7 8 15 7 7 2 16 5 11 16 2 14 16 2 14 16 16 16 16 16 16 16 16 16 16 16 [5 15 15 15 14 1 ] 16 NA 16 16 16 16 16 16 16 16 3 13 16 2 14 16 2 2 212 #4 1 16 16 16 11 4 1 16 16 16 16 16 9 5 1 15 16 16 16 16 1 #113 15 16 16 16 16 15 15
Atelerix albiventris
(N = 25; M-11, F-14;
J-4, MA-13, W-8)
NO. 3340
Atelerix algirus (N = 21; M-7, F-13, ?-1;
J-5, MA-7, W-9)
0
23 23
21
23 25
25 14
it 10
24 24 24
25 25
25 25 13
20
25 25 19
18
1
4 20
22 22
22 17
25
25
10 21 21 22 24
It 13 2t 23
25
o- 2 4 RP lo 7 tS BAL? 5| 15 15
20 ig 12
NA NA
NA NA
23121 21
24| 9 11 20
42 my 21
49. a 2]
NA NA
Na | 21 2]
oy |S 24
22! 3 18 2]
19/20 1 2] 15 22} 20 ] 21 19 19 21 21 23 |e 36 21
25/21 2
24 25 16 1 17 25 20 1 vy
25 25 21 21 25 | 21 21
19/21 2
25 41 21
21! 6 8 14
21121 2
92} 119 1 24
22111 3 14
22} 9 10 19
24 21 yy
NA NA
NA NA
NA NA
24/21 oy
24 | 21 24
24/21 pa
NA NA
NA 1 2]
25 | 21 21
25 | 21 21
NA NA
25 21 21
25/19 1 20
24 | 21 21
13 | 21 21
PS AT oF 1.31
23 21 1
22112 7 1 20
25 21 2]
NA NA
NA NA
25 | 21 2
25 | 21 21
| 25] 12 7 1 20
7 24) 8 5 7 20 25/17 4 2)
25| 1 18 19
2001 GOULD: DENTAL MORPHOLOGY OF HEDGEHOGS 39
APPENDIX 3 Continued
Hylomys sinensis (N = 25; M-15, F-9, ?-1; J-0, MA-24, W-1)
Echinosorex gymnura (N = 32; M-14, F-16, ?-2; J-7, MA-18, W-7)
Aylomys suillus Atelerix albiventris (N = 16; M-S, F-11; (N = 25; M-11, F-14, J-3, MA-11, W-2) J-4, MA-13, W-8)
Atelerix algirus (N = 21; M-7, F-13, 7-1; J-5, MA-7, W-9)
TS# edo 3 Ay Oe tt 2 54 04 = Asier 169 I 1 170 25 5 15 20 171 15 15| 1 24 25|12 9 21 172 | 32 32 16 16| 24 24) 21 21 173 | 32 32 167 2 16 NA NA 174 [31 3] 115 16 NA NA 175 28 3 31 32 10 15 NA NA 176 | 32 32 16 16 NA NA 177 32 32 15 15 NA NA 178 3h 3} 8 6 1 15 NA NA 179} 5 15 10 30 16 16 NA NA 180! 31 3] 15 1s) 25 25: i 21 181 | 31 31 15 15| 25 25] 21 21 182 | 5 20 2 at 2 16/19 3 22/19 2 21 183 | 13 16 1 30; 25 5c = Ha 14) 25 25'|, 24 21 184; 22 5 3 1 31 | 25 251 6 9 1 16) 25 25/19 2 21 185 | 29 29 na| 16 16) 25 25) 20 20 186 | 6 24 30 na] 8 7 1 16/25 25)19 1 20 187| 2 26 1 29 na] 9 6 1 16| 22 25/17 3 20 188 | 31 31 na | 16 16] 25 25| 2 19 21 189 | 30 30 wa] 9 9/11 M1 NA 190} 25 3 28| 25 25| 7 6 2 15] 9 13 224; 19 19 191] 29 1 30 NA| 9 9111 i NA 192| 3 24 1 28} 2 23 25} 8 6 2 16) 8 13 1 22} 1 20 21 193 4 41 2 2110 1uls 1 9) 1 194] 12 15 3 131; 23 2 25 15 1 16 25 25| 3 117 21 195 31 31) 25 25| 16 16| 25 25) «42 21 196 | 31 31 | 25 25 | 16 16 | 25 25/19 2 21 197 32 32/16 5 21 15 1s| 24 mm) 5 7 3 15 198 na | 25 251 15 15 | 24 24 | 21 21 1991 4 12 2 18/24 1 25; 15 15| 13 5 18] 21 21 200 32 32) 25 25/6 8 1 530 2-48 1 21 NA 201 | 11 10 4 25/19 3 224° 5 6 5 16|19 1 20} 21 21 202130 1 31] 10 10 323/13 2 15 | 20 20 | 21 21 203 | 30 1 31 | 25 25] 15 15 | 23 23 | 21 21 204| 5 27 32/13 11 125) 16 16 | 25 25) 21 21 205 32 32| 25 25| 16 16| 25 25) 21 21 206 | 32 32| 25 25) 15 15 na} 21 21 207 | 32 32 | 25 25115 15 | 21 20h] 21 208 | 32 32| 25 25/15 15 | 23 23) 21 21 209 | 32 32 | 25 25 [15 is | 1 8 9 21 21 210} 31 31 25 25 15 15 23 23 21 21 211; 15 10 1 26 25 25| 15 15 23 23) 21 21 212 6 26 32.) 35 25; 15 15; 24 24} 2 18 121 213 | 15 10 3 28/18 6 | PD. 59 16| 212 5 1 20:3 3 9 217 214} 3] 31| 23 23 NA} 17 2 19 NA 215 | 31 31| 6 15 425] 16 16; 22 22) 34 2] 216|21 9 30|20 3 225] 16 16; 121 3 251 20 1 21 217; 31 31| 25 25| 15 15! 4 13 4 21/12 8 1 21 218| 1 22 4 27 25 25 16 16 21 21+ 1 112 3°17 219 | 32 32} 25 25 | 16 16/22 | 23 | 21 21 220) 32 32124 24] 15 15 | 21 21] 4 8 12 221 32 32/24 24} 16 16| 20 20/19 1 20 222 | 32 32 | 25 25 | 16 16 | 25 25 | 21 21 223 31 31) 25 25} 16 16; 25 25; 21 21 224] 31 31/10 4 8 22; 16 16 8 1 920 20
40
AMERICAN MUSEUM NOVITATES
NO. 3340
APPENDIX 3 Continued - Echinosorex gymnura Hylomys sinensis Hylomys suillus Atelerix albiventris Atelerix aigirus (N = 32; M-14, F-16, 9-2; | CN = 25; M-15, F-9, ?-1,; (N = 16, M-5, F-11, (N = 25, M-11, F-14; (N = 21; M-7, F-13, 9-1; J-7, MA-18, W-7) J-0, MA-24, W-1) J-3, MA-11, W-2) J-4, MA-13, W-8) J-5, MA-7, W-9) TS# 0 12 3 4AT!|] 0 12 3 4 AT 225 25 25 21 21 226 25 25 | 21 2] 227 25 25) 6 10 16 228 10 2 9 1 22 | 21 2] 229 25 25 | 21 21 230 14 1 1 16/10 5 1 16 231 Diet 2 14) 6 5 i 232 9.9 2 20) 6 14 1 21 233 NA NA 234 NA NA 235 25 25 21 pal 236 1 2 1 4/21 2] 237 25 25 21 2] 238 25 25 21 2] 239 25 25| 1 19 I 2] 240 22 22 | 21 2) 241 20 25 21 2] 242 25 25 2] 21 243 25 25 | 21 2] 244 25 25 21 2] 245 25 25 | 21 21 946 25 25 16 117 947 25 25 2] 21 248 NA 21 21 249 25 25 20 20 Frinaceus amurensts Erinaceus europaeus Hemiechinus aethiopicus Hemiechinus auritus Hemiechinus hypomelas (N = 11; M-3, F-6, 2-2; + (N= 24; M-10, F-8, ?-6; | (N = 25; M-13, F-11, 2-1; | (N = 25; M-12, F-4, 7-9; (N = 19; M-10, F-9; L J-2, MA-7, W-2) J-2, MA-15, W-7) J-3, MA-15, W-7) J-8, MA-11, W-6) J-4, MA-11, W-4)
TS#; 0 1 23 4 AT;,O 12 3 4 AT/]O 1 23 4 AT) O01 23 4 AT} 0 12 3 4 AT 24/25 25 | 25 25 | 19 19
24 25 25 25 25 19 19
24 | 24 1 25 | 25 25 | 18 1 19
23 22 22 25 25! 1 4 10 1 16
24 25 25 25 25 19 19
2 22114 10 1 25; 4 17 21} 217 19
1 24; 25 25 | 25 25 | 19 19
2 23 17 6 2 25 20 3 1 24 217 19
24 | 25 R15 5 20/18 18
24) 11 14 25! 3 19 22)17 2 19
24) 1 23 24) 1 17 18 18 18
22) 17 6 23/19 J 20 | 13 13
24 23 23 22) 23 18 18
22} 8 6 3 3 20) 1 8 6 217; 2 4 9 217
24 23.2 25 24 24 19 19
5 24)22 1 1 24 | 16 | 17 | 19 19
24/19 6 25 24 24 19 19
] 23] 2 23 25) 5 11 16 19 19
23 25 25 16 16 18 18
24) 1°11 13 25 15 15| 1 2 14 1 18
24 25 25 24 24 19 19
NA NA NA NA
NA NA NA NA
NA NA NA NA
2001
31 32 33 34 35 36 37 38 39 40 4l 42 43 44 45 46 47 48 49 50 51
52 |
33 54 55 56 57 58 59
60 |
6] 62 63 64 65 66 67 68 69 70 71 72 73 74 75 76 77 78 79 80 81 82 83
FErinaceus amurensis
(N = I1; M-3, F-6, 9-2;
woo RUA
10
10
11 II It ik II
1! 1] 1]
1] ia 1]
J-2, MA-7, W-2) 23 4AT
I]
10
10
11 I
— ma AD
1i
11 1]
10
11
GOULD: DENTAL MORPHOLOGY OF HEDGEHOGS
Erinaceus europaeus (N = 24; M-10, F-8, ?-6; J-2, MA-15, W-7)
APPENDIX 3 Continued
Hemiechinus aethiopicus (N = 25; M-13, F-11, 7-1;
J-3, MA-15, W-7)
Aemiechinus auritus
(N = 25; M-12, F-4, 2-9;
J-8, MA-I1, W-6)
10 1] 10 10 10 10 10
10
10 10 10 NA 10 10 10 10 NA 1] 11 11 It it 1] 1] I! lt 1] 1] 11 11 11 11 i] i i 1] 1 1] 11 11 II I} 8) iD 1] 1] 1] 1] 1]
0
23 19 14
12 aI
22 22
20
13
24 24
24 24
24
18
17
24
13
22
24 21
24 24
]
bo tA
22:
16
22
22
22
24
24
24
24
22 13 20
24 24
25 24 19 22 21 21 21 21 NA 2) 2] NA NA 20 21 21 NA 21 21 NA 21 NA 2] 25 25 25 25 25 23 25 24 a4 25 24 24 25 25 25 25 25 25 25 22 17
2 13
20 at 24 24 24 25 25 25
23 4AT/O123 4 AT 25 4 3 18 6 222} 4 4 14 24/13 2 8 21 221) 119 1 21 1 22 1 20 1 22) 2] 118] 21 13 27\) 2) fet%| 92 11 3 16 16 22 | 20 22| 9 12 22| 21 NA 22|| 2 20 22 21 1 21 22/ 8 13 NA 72) Pl s2G 24| 25 24 | 25 24 | 25 24 | 25 24 | 25 24) 23 24} 4 21 24 | 24 24 24| 24 24; 25 19 1 22) 4 10 10 1 23) 320 1 1 23) 218 5 23 25 24) 25 24 | 25 24 24 25 5 223} 8 li 4 24| 25 22/21 1 22 oi 216) 7 2 2 2 22 | ble 8 24 24 1 20 24 | 24 i 1 23] 24 9 6 23| 24 24| 25 24 | 25 24 | 25 24/19 1 l
21
0
Oo ta
25
16
23
I
25 20
25 25 25 25 25
25 15
| 25
25
11
25
23
25 25
25 25
1
25 25 10
18
23
16
20
21 25
25
25
—
25 25
Ze A ae ON
21
14
25
25
23 25
T
41
Hemiechinus hypomelas
0
ho lo
17 16
19 19 7 18
17 17
1
_
Nm ~
17 15 17
10 17
19
16 12 19 18
17 17
11
(N = 19; M-10, F-9; J-4, MA-11, W-4)
23 4 A
42 AMERICAN MUSEUM NOVITATES NO. 3340
APPENDIX 3 Continued
Erinaceus amurensis Erinaceus europaeus Hemiechinus aethiopicus Hemiechinus auritus Hemiechinus hypomelas
(N = 11; M-3, F-6, 7-2; | (N = 24; M-10, F-8, 9-6, | (N = 25; M-13, F-11, ?-1,; |; (N = 25, M-12, F-4, ?-9; (N = 19; M-10, F-9;
J-2, MA-7, W-2) J-2, MA-15, W-7) J-3, MA-15, W-7) J-8, MA-11, W-6) J-4, MA-11, W-4) O° To? Bet A Toe Ol 23 46 AT | 2) 23 4 A TE Od 2 oe 4a AT 19 5 24| 9 7 8 24; | 14 ft "22 17 17 24 24 25 23 25 23: Be5 2 15 24 24 25 25 1 24 25 17 17 24 24 25 25 25 25 17 17 24 24 21 21 25 25 TF 17 22 22 18 ] 19) 1 1F 18 15 15 NA 1 1) ul | NA 24 24,17 2 R204. 25 25 | 16 16 13k 1 17: tl 6 2 19 | 25 25 | 15 15 1 16; 4 12 2 18 | 25 29 )|..2. 7 4 14 24 24 | 25 25-7 25 ao 4l2 17 24 24 | 23 23 | 25 25 17 17 24 24 | 25 25 | 25 25.) 17 17 22 Dealer 23 25 25 17 17 15 6 3 24/21 2 1 24) 25 25 | 17 17 24 24 | 24 24 | 25 25 | 17 17 100 | 1 10 Ww! 517 224) 24 1 255)" “25 251 15 15 101; 6 2 3 1:21 1 1 } 24) 222 1 25 18 4 22| 1 14 15 102/11 11) 24 24) 7 17 24,17 4 122; 8 7 15 103 | 11 11 | 24 24 | 23 23:18 3 21; 15 1 16 104; 9 2 11 j 24 24) 8 25 1 24] 22 22 1 15 105 a 11 24 24 25 25 25 25 19 19 106 | 11 1] | 24 24 | 24 24 | 23 23 NA 107 | 11 1] | 22 1 1 24 | 25 25} 8 2 12 22,17 1 1 19 108 11 1 24 24 25 Zot lA 22; 1 18 19 109 1] 1 24 24 25 25 25 25 | 19 19 110 1] il oo earl 23 25 25 25 25 19 19 111 Il tH 24 24] 6 12 2 20 25 25 16 16 112} 11 11}16 5 21/16 4 20} 21 2 1 24] 15 1 16 113 11 H 16 16 1s 15 2] 21; 1 14 15 114 11 11 16 16 25 25} 1 18 19 16 16 115 NA NA NA NA NA 116 NA NA NA NA NA 117 | il 1] | 24 24 | 25 2525 25 | 19 19 118; 3 8 1} /} 18 6 24 | 25 25 | 25 25 | 19 19 119 iF ie 24 24 25 25 25 25 19 19 120 11 1 24 24 os) 25 25 25) 17 1 18 12] NA NA NA NA | I 122 NA NA NA NA NA 123 | 11 11 | 24 24/24 | 25; 1 24 25) 16 16 124]; 5 6 M200 3 ll 24; | 23 24; 1 21 1 23) 5 10 3 18 125]; 8 2 10|16 7 23 14 10 24) 8 13 21/9 3. 3 17 126| 6 1 4 11/17 7 24; 5 5 12 2 24/10 1] 21; 113 4 18 127; 8 3 11/17 5 22 3 2-18 23| 6 15 21) 4 1°13 18 128) 9 2 11} 12 12 24/19 3 22;14 7 21119 19 129 | 11 1] | 24 24) 2b | 22 | 25 25 | 19 19 130 fies 10 8 14 1 23 6 5 i] 25 25 2 2 4 131 1} 1h 24 24 24 24 25 25 19 19 132 il MN 24 24 24 24 25 5 19 19 133} 11 1] | 24 24 24 24 | 25 25) 19 19 134 | 11 1 24 24 24 24 25 25 19 19 135 11 11 24 24 24 24 25 25 19 19 136; 3 8 11} 7 12 19:21 21) 7 18 25/10 1 6 [7 137 | 11 11} 18 1 19/15 4 1%)-LB- *2 20/16 17 138 I 1] 24 24) 1 22 23 25 25 19 19
139} 5 1 5 W)15 7 2 24; 6 14 20 24 J 25 19 19
2001 GOULD: DENTAL MORPHOLOGY OF HEDGEHOGS 43
APPENDIX 3 Continued
Erinaceus amurensts Erinaceus europaeus Hemiechinus aethiopicus Hemiechinus auritus Hemiechinus hypomelas
(N = 11; M-3, F-6, ?-2; GN = 24; M-10, F-8, 7-6; | (N = 25; M-13, F-11, 7-1; | (N = 25; M-12, F-4, 7-9, {N = 19; M-10, F-9;
J-2, MA-7, W-2) J-2, MA-15, W-7) J-3, MA-15, W-7) J-8, MA-11, W-6) J-4, MA-11, W-4) oO 123 4AT]0O 123 4 AT) 0123 4 AT 3 17 20 25 25| 4 13 1 18 25 25 25 25 19 19 NA NA NA | Na | NA | NA 144 | NA NA NA NA NA 145 | 1 1E | 24 24 | 25 25 | 25 25) 19 19 146 | 11 li | 24 24 | 25 25 | 25 25 | 18 18 147 | 11 11 | 24 24 | 25 25 | 25 25) 19 19 148 | 11 11 24 24 NA NA NA 149 11 11 24 24 25 25 25 25 19 19 150] 11 11 | 24 24 | 25 25 | 25 25119 19 151 11 11 | 24 24 | 25 25 | 25 25119 19 152 NA NA NA NA NA 153 | 1 1 24 24 NA 25 25 | 19 19 154] 7 4 11 | 24 24/19 3 2 24125 25|14 3 2 19 155 | 10 10 | 24 24/12 6 18; + 24 25) 4 7 11 156 | 10 10 | 23 23; 1211 12) {| | | 222 4 157; 5 3 2 10 231 24) 1 19 2, 22 25 25 13 13 158 10 10 24 24| 3 18 21 25 25 13 13 1I59| 5 3 8 24 24) 2 18 4 24 25 25 13 13 160 Il 1] 24 24) 1 24 25 | 25 25 19 19 161 NA NA NA NA NA 162 NA NA NA NA NA 163 | 11 11 | 24 24 | 25 25 | 25 25 | 17 1 18 #64 | 11 1] | 24 24/23 2 25 | 24 24119 19 165 | 5 4 2 11} 22 I 23)13 2 9 24 24 24/12 2 1 | 16 166) 9 1 10 16 8 24 21 2 23 24 24 | 10 5 15 167) 5 6 11) 5 19 24] 9 13 3 25 24 24] 6 13 19 168 | 10 10 | 22 22| 3 18 1 22 25 25] 1 14 15 169 | 10 10 | 22 22]; 1 3 4 NA 1 ] 170; 2 8 1 11] 3 21 24] 2 20 22 25 251 5 10 15 171) 11 11| 3 21 24 23 23 25 25 15 15 172 It 11 24 24 25 25 25 25 19 19 173 NA NA NA NA NA 174 NA NA NA NA NA 175 NA NA NA NA NA 176 NA NA NA NA NA 177 NA NA NA NA NA 178 NA NA NA NA NA 179 NA NA NA NA NA 180 1] 11 24 24 24 24 25 25 19 19 181 1] li 24 24) 10 12 22 25 25 19 19 182 | 11 115 24 24) 1 22 23 | 25 25 | 18 18 183 1 1 24 24 24 24 25 25 19 19 184 | 1] 11 | 24 24) 23 I 24 | 25 25 | 18 ] 19 185 li 11 24 24 23 23 25 25 18 18 186 | 11 1] | 24 24 | 23 23 | 25 25 18 18 187 11 11 | 24 24/17 4 21 | 25 25 | 16 16 188 | 11 11 24 24| 3 20 1 24 25 25] 1 18 19 89 | U1 11 | 24 24; 8 8 NA] 9 9 190 | 11 ll] 6 16 1 23] 7 17 24 25 25) 9 18 191] 11 EY] 7 7| 8 8 NA) 7 3 10 192 11 1}; 1 23 24] 2 19 2 23 2] 21; 5 9 3°17 193 NA| 1 1) 1 4 5 NA| 6 l 1 8 194 5 6 i] 1 23 24) 3 21 24 25 25 19 19
195 1! 1] 24 24 24 24 25 25 19 19
4 AMERICAN MUSEUM NOVITATES NO. 3340
APPENDIX 3 Continued
Aemiechinus auritus (N = 25; M-12, F-4, 7-9;
Erinaceus amurensis (N = 11; M-3, F-6, 7-2;
Frinaceus europaeus (N = 24; M-10, F-8, 7-6;
Hemiechinus aethiopicus (N = 25; M-13, F-11, 7-1;
Hemiechinus hypomelas (N = 19; M-10, F-9;
J-2, MA-7, W-2) J-2, MA-15, W-7) J-3, MA-15, W-7) J-8, MA-11, W-6) J-4, MA-11, W-4)
0123 4AT
19 19
14 2 16
18 18
199 | 11 11 | 24 24 | 25 25 | 25 25419 19 200 | 11 11 | 24 24 25 25 25 25 16 16 201} 11 Ht] 8 11 1 20] 25 25113 4 17/13 3 2 18 202 1 11 | 24 24 25 25 | 25 25/18 18 203 | 11 1 | 24 24| 713 1 1 22 25 251/13 3 16 204} 11 1 | 24 24 25 25} 4 4 19 19 19 205 ml i 24 24 25 25 25 25 19 19 206 | 11 1 24 24 25 25 25 25 19 19 207 | 11 11 | 24 24 | 25 25 | 25 25119 19 208 1 1 24 24 | 21 21 25 25 18 18 209 11 Hl 24 24 19 19 25 25 17 17 210 1] 7 24 24 24 24 25 25 19 19 21 11 i 1 23 24 15 6 21 25 25 19 19 212 11 1 24 24| 1 24 25 1 24 25 19 19 213 10 10; 1 9 6 420114 1 7 9. 24 | Paty 4 18/13 3 117 214 if 1 24 24 5 7 12 7 1 8 18 18 215 il 11] 1 23 24 14 14 9 9 5 5 216| 2 9 ist 24 24 16 9 25 9 12 21 4 13 17 217 1 1 24 24| 123 1 25 25 25 Mes 1 18 218 10 111 21 21] 216 4 2 24/1 16 118! 1 213 2 18 219/11 11 | 24 24 | 25 25 | 25 25/19 19 220; 7 4 1} 1l 6 17/14 5 1 20 25 25| 6 6 1 13 2211 11 11 24 24 25 25 | 25 25 15 15 222 11 11 | 24 24 | 25 25 | 25 25/19 19 223 1 11 24 24 25 25 25 25 19 19 224 11 11 24 24 25 25/24 1 25 5 5 225 11 11 24 24 25 25 25 25 19 19 226 | 11 11} 24 24 | 25 25} 25 25119 19 227| 9 19 24 24) 1 21 22| 16 16 228 | 11 24) 25 25) 25 25/19 19 2291 11 24 | 25 25 | 25 25119 19 2301 8 24| 7 14 21) 510 1 16/7 9 16 231} 1 4| 1 4 5| 1 3 Al 5 1 6 232 24) 219 1 22] 1 24 25 18 18 233 NA NA NA NA 234 NA NA NA NA 235 23 24 24 | 25 25/ 1 5 10 16 236 24 21 pil 25 25 16 16 237 24 24 24 25 25 19 19 238 24 24 14 25 25 19 19 239 24 23 1 24 25 25 19 19 240 24 | 24 24 | 25 25} 19 19 241 24 19 19 25 25 19 19 242 24) 21011 1 24 8 12 20 5 12 7 243 24 | 24 24 | 25 95119 19 244 24 20 20 25 25 19 19 245 24 | 20 20 | 25 25| 19 19 246 | 24 20 20 25 25 NA 247 24 20 20 25 25 19 19 248 23 na | 25 25 NA 249 | 22 16 16 25 25 NA
2001 GOULD: DENTAL MORPHOLOGY OF HEDGEHOGS 45
APPENDIX 4
Transformation Series Recovered for Phylogenetic Analysis Abbreviations: PA# = number assigned to the transformation series for the phylogenetic analysis; TS# = transfor- mation series number; ECHG = Echinosorex gymnura; HYLU = Aylomys suillus; ATXA = Atelerix albiventris; ATXG = A. algirus; ERIA = Erinaceus amurensis; ERIAE = E. europaeus; HEME = Hemiechinus aethiopicus; HEMA = #4. auritus; HEMH = H. hypomelas.
Taxon PAH TS# Transformation series ECHG HLYS HYLU ATXA ATXG ERIA ERIAE HEME HEMA HEMH l 4 I1: (0) present; (1) absent 0/1 0 0 0 0 0 0 0 0 0 2 5 I], size: (QO) normal; (1) enlarged (> 12) | 0 I | 1 | 1 1 l | 3 6 12; (0) present; (1) absent 0 0 0 0 0 0/1 0 O/1 0 0/1 4 8 I2, size: (0) > 13; 1) = 13; @) <3 0 0 o/1 2 2 2 2 2? 2 2 5 10 13; (0) present; (1) absent 0 0 0 0 0 0 O/1 0 0 0 6 24 Pl: (0) present; (1) absent 0 I 0/1 1 1 O/1 1 i 1 1 Py “27 P1, roots: (0) one; (1) two 0 NA 0/1 NA NA 0 NA NA NA NA 8 28 P2: (Q) present; (1) absent 0 0 0 0 0 0 O/1 0 0 0 9 32 P3: (0) present; (1) absent 0 0 0 0/1 0 0 0/1 O/1 0 0 10 39 P3: (0) normal; (1) reduced 0 ] 1 1 1 0 I 1 0/1 0/1 1] 42 P3, protocone: (0) !/2 the height of the NA NA NA ] 1 ! i NA 0/1 ] paracone; (1) much smaller 12 43 P3, paracone shape: (0) conical; 0 0 0 O/1/2 0 | 0 0 0 0 (1) crescentic 13. 46 P3, centrocrista (paracone-metacone): 1 1 ] j NA NA NA NA NA I (0) present; (1) absent 14 47 P3, hypocone: (0) present; O/1/2 1 1 1 1 1 1 1 I 1 (1) vestigial or absent 15 54 P4, hypocone: (0) present; (1) absent 0 0 0 0/1 0 0 0 0 0 0/1 16 55 P4, hypocone height: (0) < protecone; 0 0 0 0 0 0 0 0 0 0 (1) = protocone 17 56 P4, hypocone gross size: (0) < protocone; 0 0 1 of 0 0 1 0 0 1 (1) = protocone 18 357 P4, protocone position (to paracone): 0 8) 0 0/1 0 0 0 0 0 0 (0) anterior; (1) posterior 19 67 P4/M1 position: (0) oblique to tooth row; ] 1 ] 1 1 1 1 1 ] ! (1) not 20 «68 M1, size: (0) largest tooth; (1) not 0 0 0 0 0 0 0 0 0 0 2) 69 MI, shape: (0) T-rectangle; (1) A/P-rect- 2 2 1/2 2 2 2 0, 2 2 1/2 angle; (2) quadrate 22 «7 M1, lingual roots: (Q) separate; (1) fused 0 0 NA 1 i 1 I 1 1 NA 23 0672 M1, metaconule: (0) present; (1) absent 0 0 0 0/1 0 0 0 O/T 0 0 24 «77 M1, protocone shape: (0) crescentic; 0 0 0 0 0 0 0 0 0 0 (1) conical 25 78 M1, protocone position (paracone): 0/1 0 0 0/2 0 0 0 0 0 0 (0) anterior; (1) posterior; (2) equivalent 26 ©=««88 M1, hypocone height: (0) tall; (1) short; O/1 2 1 1 ] 1 1 ] 1 1 (2) approximately equal 27) = 93 M1, hypocone: (0) isolated; (1) attached o/1 1 1 1 1 1 } 0/1 1 i to other crests/cusps 28 94 M1, metastyle: (0) present; (1} absent 0 0 0 0 0 0 0 0 0 0 29 «OS M1], metastyle apex (cone itself): (0) high; ] 0 | 1 1 0 0 0 0 ] (1) low 30-96 M1, metastyle position (relative to meta- 0 0 0 0 0 0 0 0 0 0 cone): (0) labial; (1) posterior 31 99 M1, metacrista: (0) present; (1) absent 0 0 0 0 0 0 0 0 0 0 32 105 M2, shape: (0) T-rectangle; (1) A/P-rect- 2 2 2 2 2 2 2 2 2 2 angle; (2) quadrate 33 106 M2, lingual roots: (0) fused; (1) separate 1 ] NA NA 0 0 0 0 0 NA 34 «109 M2, protocone size: (0) = paracone; 1 1/2 1 NA } 2 2 1 2 0 (1) larger; (2) smaller 35 III M2, hypocone: (0) isolated; (1) not 0/1 I l ] ] ] l 0/1 1 j
3606117 M2, metastyle: (0) present; (1) absent 0 0 0 0 0 0 0 0 0 0
46 AMERICAN MUSEUM NOVITATES NO. 3340
APPENDIX 4 Continued Taxon PA# TS# Transformation series ECHG HLYS HYLU ATXA ATXG ERIA ERIAE HEME HEMA HEMH 37) 121 M2, posthypocrista: (0) to postcingulum, NA 0 NA NA NA NA NA NA NA NA (1) not 38 129 M3: (0) present; (1) absent 0 0 0 0 0 0 0 0 0 0 39 «6133 M3, protocone size: (0) large; (1) small 0 0 ] 0 0 0 0 1 0 0 40 134 M3, main cusps: (0} equally developed; ] 0 0 0/1 0 0 1 1 I | (1) not 41 135 M3, metaconule: (0) present; (1) absent 0/] | l 1 ] I ] 1 1 ] 42 141 il: (0) present; (1) absent 0 0/1 0 } ] I 1 | 1 1 43 142 il, size: (0) =12; (1) > 12 0 1 ] NA NA NA NA NA NA NA 44 143 il, shape: (0) spatulate; (1) conical 0 0 0 NA NA NA NA NA NA NA 45 144 il, root: (0) short; (1) long I 1 ] NA NA NA NA NA NA NA 46 145 i2: (0) present; (1) absent 0 0 0 0 0 0 0 0 0 0 47 146 i2: (0) enlarged; (1) reduced 0 1 0 0 0 0 0 0 0 0 48 147 i2, shape: (0) spatulate; (1) conical 0 0 0 0 0 0 0 0 0 0 49 149 i2, anterior midline crest: (0) ends pos- 2 0/1 NA NA 1 ] I 1 2 l terior to protoconid; (1) not $0 «150 i3; (0} present; (1) absent 0 0 0 0 0 0 0 0 0 0 51 151 i3, size (relative to i1/i2): (0) smaller; 0 0/2 0 0 0 l 0 0 0 0 (1) equal; (2) larger 52 152 lower canine, size: (0) = pl; (1) > pl 1 NA 1 NA NA NA NA NA NA NA 53 153 lower canine, morphology: (0) like 12/p2; 1 0/1 I 0 1 I 1 1 1 0 (1) net 54160 pl: (0) present; (1) absent 0/1 1 0 1 1 l 1 0/1 0 I 55 16] pl, roots: (0) single; (1) partially divided 0 NA 0 NA NA NA NA NA NA NA 56 163 p2: (0) present; (1) absent 0/1 0 0 0 Q 0 0 0 0 O/] S57 164 p2, roots: (0) one; (1) two 0 0 0 0 0 0 0 0/1 0 0 58 172 p3: (0) present; (1) absent 0 0 0 ] 1 | } 1 1 l 59 173 p3, roots: (0} two; (1) one; (2) two fused 0 I Of1/2 NA NA NA NA NA NA NA 60 175 p3, cusps: (0) two; (1) one; (2) three 1 l oO/1/2 NA NA NA NA NA NA NA 61 176 p3, posterior margin: (0) wide; (1) narrow 0 ! ] NA NA NA NA NA NA NA 62 177 p3, metaconid crest: (0) present; (1) absent NA 0/1 ] NA NA NA NA NA NA NA 63 178 p3, posterolingual cusp: (0) prominent, 1 1 Of) NA NA NA NA NA NA NA (1) weak/absent 64 179 p3, cingulum: (0) present; (1) absent; O/1/2 ] ] NA NA NA NA NA NA NA (2) partial 65 180 p4, talonid: (0) elongated; (1) short 1 1 1 l i 1 1 ] 1 | 66 181 p4, talonid: (0) greatest breadth of tooth; 0 ] 0 1 ] 1 ] 0/1 1 1 (1) not 67 183 p4, posterolabial cuspule to protoconid: Of] ] ] 1 I I 1 1 1 1 (0) present; (1) absent 68 [85 p4, paraconid height: (0) = protoconid; 1 NA 1 1 1 1 1 1 ] 1 (1) < protoconid 69 189 p4, protoconid size: (0) > metaconid; 0 NA 0 0 NA 0 0 0 NA 0 (1} not 70 191 p4, metaconid size: (0) small; (1) large 0/1 NA 0 0 NA 0 0 0 NA O/1 71 195 p4, size: (0) = ml; (1) smaller 1 ] ] 1 1 ] ] ] l 1 72 196 prevallid shear: (0) present; (1) absent 0 0 0 0 0/1 ] 0 0 0 0 73 198 mil, trigonids: (0) high; (1) low NA 0 I 0 0 0 0 ] 0 1 74 «205 ml, lingual wall: (0) markedly concave; 1 1 ] ] l ] 1 1 1 1 (1) not 75 206 mi, talonid: (0) enclosed lingually by 1 1 1 NA 1 0 1 1 1 ] entocristid; (1) not 76 207 ml, talonid: (0) opens posteriorly; 0 0 0 0 0 0 0 0 0 0 (1) closed 77 208 ml, hypoconid: (0) isolated; (1) not 0 l 0 0 1 1 1 0 l l 78 209 ml, entoconid size: (0) > hypoconid; 2 0 0 0/3 2 3 3 0 3 3
(1) > paraconid; (2) = cusps; (3) > than both
2001
GOULD: DENTAL MORPHOLOGY OF HEDGEHOGS
PA#
79 80
$1
82
83
Bd 85
86
87
88
89
90
9}
92
93
94
95
96 97
98
99 100
TS#
210 219
220
222
223
225 226
228
229
233
234
235
236
237
238
240
241
243 244
245
247 248
APPENDIX 4
Continued
Transformation series
ml], entostylid: (0) present; (1) absent
ml, paraconid to protoconid: (0) lingual; (1) labial; (2) aligned
ml, paracristid: (0) terminates at mid- anterior; (]) not
m2, size: (0) shorter than m1; (1) larger than mi
m2 trigonids: (0) high, short talonid; (1) low, expanded talonid
m2, paraconid: (0) present; (1) absent
m2, paraconid shelf: (0) present; (1) absent
m2, paracristid lingual extension (to m1): (0) equal; (1) labial; (2) lingual
m2, entostylid: (0) absent; (1) weak; (2) strong
m2, paraconid to metaconid: (0) lingual; (1) labial; (2) aligned
m2, paraconid to protoconid: (0) lingual; (1) labial; (2) aligned
m2, hypoconid to protoconid: (0) lingual;
(1) labial; (2) aligned
m2, entoconid size: (0) = protoconid; (1) tallest cusp; (2) = hypoconid; (3) = metaconid
m2, talonid: (0) posteriorly narrow, (1) not
m?, talonid basin: (0) lingually enclosed;
(1) not
m?, cristid obliqua contact point: (O) base of protoconid; (1) higher
m?, entocristid: (0) high; (1) low: (2) absent
m3: (0) present; (1) absent
m3, paraconid: (0) swollen; (1) normal; (2) absent
m3, paraconid shelf: (0) present; (1) absent
m3, talonid: (0) present; (1) absent
m3, trigonids: (0) high; (1) short; (2) low expanded
Taxon
47
ECHG HLYS HYLU ATXA ATXG ERIA ERIAE HEME HEMA HEMH
1 0
1 0
NA
NA
—
—
NA
NA
] 0/1
NA
o/1/2
NA
NA
2/3
NA
] 0
NA
NA
1 0
NA
NA
NA .
NA
NA
NA
NA
—
NA
NA
Oo
—
NA
NA
O/1/2
NA
48 AMERICAN MUSEUM NOVITATES NO. 3340
APPENDIX 5 Phylogenetic Analysis of Data Set A (a) Data matrix and (b) results of analysis |. For transformation series included in the analysis (PA#), refer to appendix 4.
(a) Matrix of 100 transformation series recovered from discrete dental analysis (refer to appendix 4). All characters are polar- ized according to outgroups (PA# 16, 17, 31, 85, and 98 are unpolarized); all are unordered.
Echinosorex gymnurus {01}100000000 ?01{012}000010 2000{01}{01}{01}010 0211{01}02001 {01}000100020 O11{O1}O{O1}OOO1 O71{012}10{01}10{01} 1071100210 1011010{12}02 2110{12}02001
Hylomys sinensis 0000017001 7011000010 2000021000 021{12}100000 1{01}101010{01}0 {OZ}? {O}}1?200011 1{OL}LL119222) 1001101010 1011010291 0110202000
Hylomys sutllus OLO{OT}O{O1}{O1}001 POELOOLOLO = {12}?700011010 0271107010 1010100070 OLLOOOOO{O12}{012} LI{OP}LIO1100 1011100010 1011010771 0110102001
Atelerix albiventris OLOZOL?O{O]}1 L{OIZ}IT{OLJOLOL}{OLJ1LO = 21{01 }Of{02} 11010 027710?00{01} 1177700090 =0?7017001??) 7777111100 =1001?700{03}1{01} £9117{012}0790 = {23}110202117
Atelerix algirus 0102017001 1071000010 2100011010 0201107000 1127900010 071170019? 9799111179 1{OL}OI101210 {01}011010772 0110102011
Erinaceus amurensis 01{01}20{01}0000 1171000010 2100011000 0202107000 1177700010 171190017? 2777111100 1101001310 0011000772 3100102011
Frinaceus europaeus O1O2{O1}127{O1}{O1}1 1071001010 2100011000 0202107001 1177900010 0711700127
2277111100 1001101310 1011010792 0110202{01}10 Hemiechinus aethiopicus O1{OL}ZOL70{O1}1 PO? 1LOO0010 = 21{OLJOOL{O1}O00 O201{01}07011 1477900010 OP1{O1}?0{O1}172? 27771(01}1100 1011100010 1011010771 311010201? Hemiechinus auritus 010201700{01} {01}071000010 2100011000 0202107001 1177700020 071090019? 2727111122? 1001101310 00110107790 =0110102010
Hemiechinus hypomelas _01{01}201200{01} 1011{01}O1010 {12}?00011010 0290102001 1177200010 07012{01}0122 799711110{01} 1011101310 1011010774012} 211010201
Outgroups: Tenrecoids O{01}O{O1}O1270{O1}O 1OLf197111 3217229992 2327799022 2401}{01}000{01}{01}20 {OD}P{OL}{OL}{OZH{O]}OLOL}{O]}O L{OL}{OL}{12}I1{O1}10{01} 11{01}027{02}410 1100?{02}0{12}0{23} 4229201710
Soricoids —- O{O1} {O1}1{01}01001 201119221{01} {23}2102{13}100{01}
92.0{02}{12}{O2}10{01}{O1} 10{01}0{01}01070 {O1}1110{01}0001 121¢01}111772 1110111210 21202{02}020{012} 311{01}201202
2001 GOULD: DENTAL MORPHOLOGY OF HEDGEHOGS 49
APPENDIX 5 Continued
(b) Phylogenetic Analysis 1, Tree Number |: Length = 105; consistency index (CID = 0.676; homoplasy index (HI} = 0.324; CI excluding uninformative characters = 0.634; HI excluding uninformative characters = 0.366; retention index (RI) = 0.528; rescaled consistency index (RC) = 0.357.
Hylomys sinensis Echinosorex gymnura Hylomys suillus Hemiechinus aethiopicus Atelerix algirus Erinaceus amurensis Hemiechinus auritus Erinaceus europaeus Atelerix albiventris Hemiechinus hypomelas
Apomorphy list
Branch PA# Steps Cl Change Branch PA# Steps CI Change nodel ~— node 2 5 1 1.000 O51 node7 — node 8 20 1 0.500 O51 50 1 1000 t>0 84 1 0500 0-51 98 1 0.333 2>1 98 ] 0.333 132 103 1 0.667 O=>1 nodes se Hadew 32 1 0.333 031 node2 — node 3 9 1 0500 1350 37 1 0667 2340 32 1 0.333 O=>1 56 1 1.000 1>0 52 1 0333 132 94 1 0500 052 37 | 0.500 1>0 node9 — Atelerix 80 1 0.250 1=>0 69 1 1.000 E> 0 albiventris, 10k 1 1.000 O03] node 3. > Echinosorex 3 1) 0.3330 f=0 node9 -» Hemiechinus 76 1 0333 OS] pial aoa? fe ie 0=>1 hypomelas 98 I 0.333 2-1] ‘ 1 aa x : node’ -—- Erinaceus 93 1 0.400 032 81 1 0600 052 i ge!
93 1 0400 1-2 node 7 —- Hemiechinus 52 ] 0.333 l>2 04 ] 0.500 O=2 auritus 57 ] 0.500 I1=>0 node3 —» Hylomyssuillus 20 1 0500 O=1 node 6 —> Erinaceus 13 1 0333 1=0 4? ] 0.500 G51 amurensis 15 1 1.000 O>1 node2 —> node 4 7 1 1000 032 54 1 1.000 0 1 25 1 1000 051 D5 te OO FO 78 1 1000 150
36 1 0.667 1-0 89 1 0.500 1>0
45 ] 1.000. QO>1 102 «10.500 OS 1 9 1 1000 150 ode dase nodes 6 i 0333. «120 node5 ~~» Atelerix algirus 32 Y 9333 Ol 80 1 0250 O0>51 node4 -— Hemiechinus 42 l 0.500 O51 81 1 0600 O32 | aethiopicus 43 1 0333 O0=>1 84 I 0.500 130 94 I 0.500 O0=>3 93 ] 0.400 17? nodel — Hylomys 29 j 1.000 |=? node5 —> node 6 37 «61 (0.667 «132 sinensis 62 1 1000 O=>1 $1 1 0.600 2-33 76 l 0.333 1>0 node 6 —> node 7 43 1 0333 O31 80 1 0.250 O01
93 1 0400 20 103 1 0667 I1=>0
50 AMERICAN MUSEUM NOVITATES NO. 3340
APPENDIX 6 Phylogentetic Analysis of Data Set B
The 29 discrete dental characters analyzed (a) are listed, along with the accompanying matrix (b) extracted from Gould (1995). The first apomorphy list (c) was generated from tree 1 in analysis 2a (19 extant taxa). The second apomorphy list (d) was generated from tree 1 in analysis 2b (only the 10 taxa considered in analysis 1 [see appen- dix 5] are treated in this analysis). Numbers in brackets [] refer to original character numbers in Gould (1995); TS# = transformation series number (see appendix 1); * = transformations not exactly as in Gould (1995); for trans- formation series included in the analysis (PA#), refer to appendix 4.
(a) Dental characters analyzed
1 [60] TS# 4 il: (0) present, enlarged; (1) present, small; (2) absent. 2 [61] TS# *5 i2 relative size: (0) greatly enlarged; (1) nearly equal to other incisors; (2) smaller than other incisors, 3 [62] TS# 8 12: (O) greater than [3; (1) less than or equal to I3. 4 [63] TS# 11 I3, number of roots: (0) one root; (1} two roots, separate; (2) two roots, fused. 5 [64] TS# 18 Cl size: (0) significantly larger than adjacent post-canine teeth; (1) slightly larger than post- canine teeth; (2) approximately equal in size to adjacent postcanine teeth. 6 [65] TS# 19 C1, number of roots: (0) two roots; (1) one root or two roots fused. 7 [66] TS# 20 C1, relative size: (0) equal to, or larger than I3; (1) subequal or slightly smaller than [3. 8 [67] TS# 152 cl, relative size: (0) approximately equal to, or smaller than P1; (1) significantly larger than pl. 9 [68] TS# 24 Pi: (0) present; (1) absent. 10 [69] TS# 160 pl: (0) present; (1) absent. ll [70] TS# *162 p2: (0) moderate size, two roots; (1) small, peglike, procumbent, one root; (1) absent. 1? [71] TS# 13 P2 roots: (0) two roots; (1) one root or two roots well fused; (2) absent. 13. [72] TS# 173 p3: (0) two roots present; larger in size than p2; (1) one root present, nearly equal in size to P; (2) absent. 14 [73] TS# 34 P3 lingual lobe (= protocone): (0) present, well developed; (1) vestigial or absent. 15 [74] TS# 39 P3 size; (0) normal: (1) reduced. 16 [75] TS# 52 P3 roots: (O) three roots; (1) fewer than three roots. 17 [76] TS# 47 P3 hypocone: (0) absent; (1) present. 18 [77] TS# 180 p4: (0) with an elongate talonid and talonid basin; (1) with a short, bicuspid or unicuspid heel. 19 [78] TS# 53 P4 shape, and hypocone: (0) quadrate, hypocone present; (1) triangular, hypocone absent or vestigial. 20 [79] TS# 59 P4 lingual roots: (0) one lingual root; (1) two unfused roots; (2) two lingual roots, fused. 21 [80] TS# *198 Trigonids on lower molars: (0) high (significantly taller than talonid), talonid short or vesti- gial; (1) low trigonid (nearly equal in height with talonid), talonid expanded, large. 22 [81] TS# *198 ml: (Q) trigonid moderate; (1) marked elongation of prevallid shear on ml. 23 [82] TS# 71 M1 lingual roots: (0) separate; (1) fused for most of the length. 24 [83] TS# *216/*#262 Distinct ectocingulum on labial side of M1] and M2: (0) absent; (1) present. 25 {84} TS# 129 M3: (0) present; (1) absent. 26 [85] TS# 1360 M3 roots: (0} three roots; (1) two roots. 27 «[86] TS# 131 M3 metastylar spur (referred to as a hypocone): (0) absent or weak; (1) present, well devel- oped on buccal side. 28 [87] TS# 132 M3 metacone conditions: (0) well developed; (1) small; (2) absent. 29 = {88} «=TS# 247 m3 talonid: (0) present; (1) absent.
(b) Data matrix extracted from Gould (1995)
Echinosorex gymnurus 1100000100 =1100000111 = 110100110 Podogymnura aureospinula 1110000111 1101000111 110100110 Podogymnura truei 1E10000111 1101000111 110100110 Hylomys sinensis 0100201011 1111110111 1410100110 Hylomys suillus 1100100000 1111110111 116100110 Hylomys hainanensis 1100100001 1111110111 110100110 Hemiechinus aethiopicus 2211200011 1121110112 111101021 Hemiechinus hypomelas 2211200011 1120100112 111101021
2001
(b)} Data matrix extracted from Gould (1995) (continued)
Hemiechinus micropus Hemiechinus auritus Hemiechinus collaris Mesechinus dauuricus Erinaceus amurensis Erinaceus concolor Erinaceus europaeus Atelerix frontalis Atelerix algirus Atelerix albiventris Atelerix sclateri Outgroups: Tenrecoids Soricoids
0.917; rescaled consistency index (RC) = 0.694.
Apomorphy list Branch
node l —- Echinosorex
node | — node 2
node2 — node 3 node2 — node 4
GOULD: DENTAL MORPHOLOGY OF HEDGEHOGS 51 APPENDIX 6 Continued 2211200011 1121110112 111101021 2211200011 1120100112 111101021 2241200011 1120700112 111101021 2211200011 1120100812 111101021 2212210011 1120100112 141101021 2212210011 4120100112 111101021 2212210011 £120100112 111101021 2211200011 1020100112 111101021 2211200011 1020110112 111101021 2211200011 1021110112 111101021 2211200011 1021110112 111101021 1100070010 0000000100 0?0000000{01 } 1100070010 ={12}001000100 0?G000000(01} (c) Phylogenetic Analysis 2a, Tree Number 1: Length = 41; consistency index (CI) = 0.756; homoplasy index (HI) = 0.244; CI excluding uninformative characters = 0.744; HI excluding uninformative characters = 0.256; retention index (RI) = Echinosorex gymnutra Podogymnura aureopinula 3 Podogymnura truei Hylomys suillus 6 Hylomys hainanensis 5 Hylomys sinensis Hemiechinus micropus Hemiechinus aethiopicus Hemiechinus hypomelas Hemiechinus auritus Hemiechinus collaris 1 Mesechinus dauuricus Erinaceus europaeus Erinaceus amurensis 10 12 Erinaceus concolor 9 Atelerix frontalis Atelerix aigirus 8 Afelerix albiventris Atelerix sclateri PA# Steps Cl Change Branch PA# Steps Cl Change 9 1 0500 150 node4 — node 5 3 t 0500 1350 3 1 O500 O-51 node 5 -» Hylomys sinensis 1 i 1.000 150 10 1 0500 O0>1 7 1 1.000 0O=>1 144001) (0.3330 (O41 node5 ~> node 6 5 1 1000 251 8 1 0750 O>1 9 1 0500 150 5 1 1.000 O=>2 node 6 — Aylomys suillus 10 1 0500 150 8 1 690500 10 node4 -> node7 1 1 1000 132 13 1 1.000 OQO>1 2 | 1.000 132 15 1 1000 051 4 1 1.000 oO0=>1 16 1 0500 O>1 13 I 1.000 12
52 AMERICAN MUSEUM NOVITATES NO. 3340
APPENDIX 6 Continued
(c) Phylogenetic Analysis 2a: Apomorphy lists (continued)
Branch PA# Steps CI Change Branch PA# Steps CI Change
node4 — node7 (contd. 20 1 1000 132 node 7 -— node & i2 1 0.333 1>0 23, 1 (1000 O01 node8 > node 9 14 1 6333 150
3 es 5 = ; node9 —> node 10 16 1 0500 150
28 1 1000 1=>2 node 1O— node 11 12 l 0.333 O=>1
29 1 1000 051 node 11 — = node 12 4 i 1.000 1>2
6 f 1.000 0O0=>1
(d) Phylogenetic Analysis 2b, Tree Number |: Length = 38; consistency index (C1) = 0.816; homoplasy index (HI) = 0.184; Cl excluding uninformative characters = 0.800; Hi excluding uninformative characters = 0.200; retention index (RI) = 0.897, rescaled consistency index (RC) = 0.732.
Echinosorex gymnura Hylomys suillus
Hylomys sinensis Atelerix albiventris Atelerix algirus Hemiechinus aethiopicus Hemiechinus hypomelas Hemiechinus auritus Erinaceus europaeus Erinaceus amurensis
Apomorphy list
Branch PA# Steps CI Change Branch PA# Steps CI Change node! — Echinosorex 8 1 1.000 O=>1 node4 -—> node 5 14 1 0333 10 nodel — node2 5 1 1.000 0 >1 node5 — node 6 16 1 0.500 I1>0 13 1 1.000 O=1 | node6 -» node7 4 1 1000 1=2 14 1 0333 O91 | Pe oh, Stig ee 15 1 1.000 O>1 ; 16 1 0500 O>1 ia ; > ieee algirus 12 } 0.333 1=0 node2 — node 3 5 1 1.000 1-2 ee inte Gen ie fi Ogee gh 9 1 0500 01 10 1 1.000 O>1 node3 — Aylomys sinensis 1 1 1.000 1=>0 7 1 1.000 O>1 node 3 — node 4 l 1 1.000 152 2 i 1.000 I>2 3 1 1000 O=>1 4 1 1000 O0O=>1 13 1 1000 1-2 20 i 1000 Il=>2 23 1 1000 O=>1 26 1 1000 O=>1 27 1 0500 150 28 1 1000 Il=2 29 i 1000 O=1