Two Kinds of Varmints: Preliminary Baraminological Analysis of Raccoons (Procyon lotor) and Opossums (Didelphis virginiana)

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1 OPEN ACCESS Report JCTS SERIES B Two Kinds of Varmints: Preliminary Baraminological Analysis of Raccoons (Procyon lotor) and Opossums (Didelphis virginiana) T.C. Wood Core Academy of Science, Dayton, TN Abstract The baraminology of two common North American mammals, the raccoon (Procyon lotor, family Procyonidae) and opossum (Didelphis virginiana, family Didelphidae), is explored using statistical baraminology. Based on a single character matrix of 40 craniodental characters, baraminic distance correlation (BDC) and multidimensional scaling (MDS) analyses could not conclusively identify the limits of the procyonid baramin. In contrast, three character matrices containing didelphids support the hypothesis that Didelphidae is a holobaramin. These studies highlight the importance of using multiple datasets for statistical baraminology studies. Editor: J.W. Francis Received January 18, 2012; Accepted January 30, 2014; Published July 7, 2014 Introduction Residents of eastern North America are extremely familiar with two nocturnal omnivores, raccoons (Procyon lotor) and Virginia opossums (Didelphis virginiana). From nighttime raids on garbage to roadkill carcasses, raccoons and opossums seem to be everywhere. Unlike so many other species, these animals have adapted well to human encroachment and habitat loss. For example, raccoons have experienced significant range expansion throughout the twentieth century (e.g., Kamler et al. 2003) and have even become invasive in parts of the Old World (Hayama et al. 2006). A nomadic, nonterritorial mammal, the Virginia opossum has also significantly expanded its range in recent history (Nowak 1999, p. 33). Raccoons are members of family Procyonidae, the membership of which has been somewhat controversial over the years. Nowak (1999) recognized seven genera: the ringtails (Bassariscus), the raccoons (Procyon), the coatis (Nasua), the mountain coati (Nasuella), the kinkajou (Potos), the olingos (Bassaricyon), and the red panda (Ailurus). Wozencraft (2005) accepted all but Ailurus. The enigmatic red panda has been considered a relative of bears (Ginsburg, 1982), raccoons (Slattery and O Brien 1995), or neither (Bininda-Emonds 2004; Flynn et al., 2005; Sato et al., 2009). The family Procyonidae is monophyletic according to the morphological analysis of Decker and Wozencraft (1991) and the molecular analysis of Koepfli et al. (2007). Opossums are members of family Didelphidae and order Didelphimorphia. Gardner (2005) recognized 87 species in 17 genera. Phylogenetic analyses of the family are sparse, and the relationship of the Didelphimorphia to Australian marsupials has been controversial (Palma 2003). The studies of Jansa and Voss (2000) and Voss and Jansa (2003, 2009) support the monophyly of Didelphidae, excluding the contested Chilean opossum, Dromiciops gliroides. Baraminology might offer additional insights into the classification of these two groups, and identifying new mammal baramins would expand creationist understanding of created kinds. Consequently, a preliminary baraminology analysis was conducted on Procyonidae and Didelphidae. Traditionally, interspecific hybridization has been an important criterion for identifying baramins, but hybridization records from both families are unavailable (Gray 1972). Instead, standard statistical baraminology methods, including baraminic distance correlation (BDC) and multidimensional scaling (MDS), were applied to four 2014 The author. This article is open access and distributed under a Creative Commons Attribution License, which allows unrestricted use, distribution, and reproduction in any medium as long as the original author and medium are credited. Citation: Wood Two Kinds of Varmints: Preliminary Baraminological Analysis of Raccoons (Procyon lotor) and Opossums (Didelphis virginiana). Journal of Creation Theology and Science Series B: Life Sciences 4:12-18.

2 Protoprocyon Procyon Paranasua Nasua Potos Parapotos Bassaricyonoides Bassaricyon Probassariscus Bassariscus Edaphocyon Arctonasua Chapalmalania Brachynasua Cyonasua Amphinasua Broiliana Amphictis Figure 1. Baraminic distance correlation results for the procyonid character set of Baskin (2004). Significant, positive BDC is represented by a filled box, and significant, negative BDC by an open circle. Black symbols denote bootstrap values 90%, and gray symbols denote bootstrap values <90%. Procyon morphological datasets, one for Procyonidae (Baskin 2004) and three for Didelphidae (Wroe et al. 2000; Voss and Jansa 2003, 2009). Methods Potosini Broiliana Amphictis Figure 2. Three-dimensional MDS results for the procyonid character set of Baskin (2004). For procyonids, I analyzed the character matrix of Baskin (2004), which consists of 40 craniodental characters and 18 taxa. The characters consist of 30 dental and 10 cranial characters. The taxa are all procyonids, and include the extinct taxa Amphirictis, Broiliana, Bassaricyonoides, Parapotos, Probassariscus, Edaphocyon, Arctonasua, Cyonasua, Amphinasua, Brachynasua, Chapalmalania, Protoprocyon, and Paranasua. There are no non-procyonid outgroups. For baraminic distance calculation, I used all the taxa and a character relevance cutoff of 0.75, consistent with other analyses of fossil taxa (Wood 2010). Thirtytwo characters were used to calculate baraminic distances. For didelphids, I analyzed the character matrices of Voss and Jansa (2003, 2009) and an additional dasyuromorph character matrix compiled by Wroe et al. (2000) that used some didelphids as outgroups. Voss and Jansa (2003) compiled a character set of 35 didelphid species from fifteen genera, representing a large portion of didelphid species diversity. There are no non-didelphid outgroup taxa. Their characters include 28 external morphological characters, 23 craniomandibular characters, 16 dental characters, and four karyotype characters. For baraminic distance calculation, I included all 35 taxa with a character relevance cutoff of Fifty-nine characters were used to calculate baraminic distances. Voss and Jansa s (2009) most recent character matrix consists of 51 taxa and 129 characters. The taxa include 44 didelphid species from 18 genera and seven outgroups. The outgroup taxa are from the families Caenolestidae (Caenolestes and Rhyncholestes), Dasyuridae (Murexia and Sminthopsis), Microbiotheriidae (Dromiciops), and Peramelidae (Echymipera and Perameles). The characters include 39 external morphological characters, 49 craniomandibular characters, 37 dental characters, and four karyotype characters. For this analysis, I used all taxa and a character relevance cutoff of 0.95, leaving 117 characters from which to calculate baraminic distances. Wroe et al. s (2000) dasyuromorph character matrix consisted of 24 taxa and 77 craniodental characters. The taxa consisted of 14 dasyuromorphs and 10 outgroup taxa, including five didelphids (Pucadelphys, Andinodelphys, Didelphis, Metachirus, and Lestodelphys). To calculate baraminic distances, I used all taxa with a 0.95 character relevance cutoff, leaving 66 characters. All baraminic distance calculations were performed by the BDISTMDS server ( Wood 2008a). Bootstrap values were calculated based on 100 pseudoreplicates. Additional statistical analyses were conducted in R ( Results Procyonidae. BDC results for the procyonids reveal four groups, consisting of (1) Amphictis and Broiliana, (2) the tribe Potosini (Potos, Parapotos, Bassaricyon, and Bassaricyonoides), (3) Protoprocyon, Procyon, Nasua, and Paranasua, and (4) the remaining members of tribe Procyonini (Figure 1). Within each group, members share mostly significant, positive BDC, and no members of the same group are negatively correlated. Between groups, significant, negative BDC occurs sparsely. The most frequent observation of significant, negative BDC is between the Amphictis/Broiliana group and the Procyon-containing group. Even in those comparisons, however, bootstrap values are low (14-84%). Only one taxon pair between groups evidences significant, positive BDC: Paranasua and Arctonasua, but the bootstrap value is only 50%. Overall, bootstrap support for most of these correlations is relatively low (median 80%). Only 15 taxon pairs (out of 153 possible taxon pairs) had bootstrap values >90%, and all taxon pairs with significant, negative BDC had bootstrap values <90%. The 3D MDS results reveal a diffuse line of taxa with tribe Potosini forming a side branch from the main line (Figure 2). The JCTS B: Life Sciences Volume 4:13

3 Monodelphis_emiliae Monodelphis_theresa Monodelphis_brevicaudata Monodelphis_adusta Thylamys_venustus Thylamys_pallidior Lestodelphys_halli Metachirus_nudicaudatus Marmosops_noctivagus Marmosops_incanus Marmosops_pinheiroi Marmosops_parvidens Marmosops_impavidus Marmosa_robinsoni Marmosa_mexicana Marmosa_rubra Micoureus_paraguayanus Micoureus_regina Micoureus_demerarae Marmosa_murina Marmosa_lepida Marmosa_canescens Gracilinanus_microtarsus Glironia_venusta Lutreolina_crassicaudata Philander_opossum Philander_mcilhennyi Philander_frenata Didelphis_marsupialis Didelphis_virginiana Didelphis_albiventris Chironectes_minimus Caluromysiops_irrupta Caluromys_philander Caluromys_lanatus Figure 3. Baraminic distance correlation results for the didelphid character set of Voss and Jansa (2003). Significant, positive BDC is represented by a filled box, and significant, negative BDC by an open circle. Black symbols denote bootstrap values 90%, and gray symbols denote bootstrap values <90%. line begins with Amphictis and ends with Procyon. The significant, negative BDC observed between Amphictis and Procyon is thus likely the result of being on the opposite ends of a linear array of taxa, as in the case of the equids analyzed by Cavanaugh et al. (2003). The 3D stress is moderately high (0.146), with a minimal stress of observed at five dimensions. Didelphidae. The BDC results for the earlier Voss and Jansa (2003) character matrix reveal two groups, corresponding to eight species of the tribe Didelphini and the remaining 27 didelphids (Figure 3). Within each group, most taxon pairs exhibit significant, positive BDC, except for the three species of subfamily Caluromyinae, which are negatively correlated with species of the genera Monodelphis, Thylamys, Lestodelphys, Marmosops, and Metachirus. Between the groups, 83 taxon pairs (of 216 possible taxon pairs) exhibit significant, negative BDC, and none are positively correlated. Bootstrap values for instances of significant, positive BDC are generally >90%; whereas most instances of significant, negative BDC have bootstrap values <90% The 3D MDS results reveal a roughly Y-shaped cloud of taxa, with the Caluromyinae and Didelphini forming the branches of the Y (Figure 4). The 3D stress of is slightly better than that observed in the procyonid MDS. The minimal stress of occurs in MDS at six dimensions. BDC results for the later Voss and Jansa (2009) dataset, which Caluromyinae Didelphini Figure 4. Three-dimensional MDS results for the didelphid character set of Voss and Jansa (2003). JCTS B: Life Sciences Volume 4:14

4 Sminthopsis_crassicaudata Murexia_longicaudata Dromiciops_gliroides Lutreolina_crassicaudata Philander_opossum Philander_mcilhennyi Philander_frenatus Didelphis_marsupialis Didelphis_virginiana Didelphis_albiventris Chironectes_minimus Monodelphis_theresa Monodelphis_peruviana Monodelphis_emiliae Monodelphis_brevicaudata Metachirus_nudicaudatus Hyladelphys_kalinowskii Glironia_venusta Tlacuatzin_canescens Marmosa_rubra Marmosa_robinsoni Marmosa_mexicana Marmosa_murina Marmosa_lepida Marmosa_paraguayana Marmosa_regina Marmosa_demerarae Marmosops_pinheiroi Marmosops_parvidens Marmosops_incanus Marmosops_noctivagus Marmosops_impavidus Gracilinanus_emiliae Gracilinanus_microtarsus Gracilinanus_agilis Gracilinanus_aceramarcae Cryptonanus_unduaviensis Cryptonanus_chacoensis Thylamys_pallidior Thylamys_venustus Thylamys_pusillus Thylamys_macrurus Lestodelphys_halli Chacodelphys_formosa Caluromysiops_irrupta Caluromys_philander Caluromys_lanatus Perameles_gunnii Echymipera_kalubu Rhyncholestes_raphanurus Caenolestes_fuliginosus Figure 5. Baraminic distance correlation results for the didelphid character set of Voss and Jansa (2009). Significant, positive BDC is represented by a filled box, and significant, negative BDC by an open circle. Black symbols denote bootstrap values 90%, and gray symbols denote bootstrap values <90%. includes additional characters and seven non-didelphid outgroup taxa, reveal two clear groups (Figure 5). The first group consists of the four species of the outgroup families Peramelidae and Caenolestidae. The remaining taxa of families Didelphidae, Dasyuridae, and Microbiotheriidae comprise the second group. Whereas most of the didelphid taxon pairs share significant, positive BDC, BDC between didelphids and non-didelphids is not always positive. The two dasyurid species are positively correlated with only ten of the 44 didelphids, and only one of those taxon pairs has a bootstrap value >90% (Sminthopsis crassicaudata and Lestodelphys halli). The two dasyurid species were negatively correlated with eight didelphid species, although the bootstrap values for those correlations was quite low (50-62%). In contrast, the Chilean opossum (Dromiciops gliroides) of the family Microbiotheriidae was positively correlated with all didelphids except three (Didelphis virginiana, Didelphis albiventris, and Chironectes minimus). The bootstrap values for 36 of those positive correlations was >90%. Dromiciops was also negatively correlated with the four peramelid and caenolestid outgroup species, although the bootstrap values for those correlations were only moderate (73-82%). Between the two main groups (one consisting of Peramelidae and Caenolestidae, and the other consisting of the remaining taxa), significant, negative BDC was observed for 174 of 188 Didelphini Caluromyinae Dromiciops Dasyuridae Peramelidae Caenolestidae Figure 6. Three-dimensional MDS results for the didelphid character set of Voss and Jansa (2009). Didelphids are shown in blue, and nondidelphids are shown in red. JCTS B: Life Sciences Volume 4:15

5 Figure 7. Baraminic distance correlation results for the dasyuromorph character set of Wroe et al. (2000). Significant, positive BDC is represented by a filled box, and significant, negative BDC by an open circle. Black symbols denote bootstrap values 90%, and gray symbols denote bootstrap values <90%. possible taxon pairs. The bootstrap support for these correlations was moderate (82.5%), with 63 taxon pairs having bootstrap values >90%. The 3D MDS results for the Voss and Jansa (2009) dataset reveals a fairly clear separation between the didelphids and the outgroup taxa (Figure 6). As noted in the 3D MDS results for the earlier Voss and Jansa (2003) dataset, the didelphids form a Y-shaped cloud with the Didelphini and Caluromyinae as the branches of the Y. The six dasyurid, peramelid, and caenolestid outgroup species are arranged in a very loose cloud that is roughly parallel to the cloud of didelphids. The microbiotheriid outgroup Dromiciops is definitely part of the didelphid cluster rather than the cloud of outgroup taxa. The 3D stress for this analysis is, however, quite poor (0.207), with a minimal stress of for MDS at eight dimensions. BDC results for the character set of Wroe et al. (2000), which focuses on 14 dasyuromorph taxa with five didelphids among the ten outgroup species, reveals only one distinct group, the eleven species of family Dasyuridae (Figure 7). Each dasyurid species is positively correlated with every other dasyurid species, and most dasyurids are negatively correlated with the outgroup taxa. The other three dasyuromorph species exhibit very limited BDC. The two thylacinids (Thylacinus and Badjcinus) are positively correlated with each other but not with any other taxa. Thylacinus is negatively correlated with the outgroup Perameles and the speckled dasyure Neophascogale lorentzii. The dasyuromorph Myrmecobius fasciatus (the numbat) is neither positively nor Dasyuridae Thylacinus Myrmecobius Badjcinus Figure 8. Three-dimensional MDS results for the dasyuromorph character set of Wroe et al. (2000). Dasyuromorphs are shown in blue, didelphids in green, and other outgroups in red. JCTS B: Life Sciences Volume 4:16

6 negatively correlated with any other taxa. The five didelphid species are part of a large group of outgroup taxa, which also includes the borhyaenid Mayulestes, the microbiotheriid Dromiciops, and the three peramelemorphs Yarala, Perameles, and Echymipera. Most members of this group share significant, positive BDC, but the peramelemorphs are positively correlated with only three outgroup taxa. The non-dasyuromorph outgroups, including the five didelphids, are negatively correlated with the eleven ingroup dasyurids. Bootstrap values for the BDC for the Wroe et al. (2000) character matrix are moderate (median 82%) (Figure 8). In general, taxon pairs with significant, positive BDC had better bootstrap values than those with significant, negative BDC. Of the 78 taxon pairs that were positively correlated, 63 had bootstrap values >90%. In contrast, only ten of the 71 negatively correlated taxon pairs had bootstrap values >90%. The dasyurids form a tight cluster in the 3D MDS results for the Wroe et al. (2000) character matrix. The outgroup taxa and the other dasyuromorph taxa form a much looser cloud adjacent to the dasyurid cluster. Despite significant, positive BDC between the thylacinids Badjcinus and Thylacinus, they do not appear closely clustered in the MDS results. They are separated by a baraminic distance of 0.317, which is only slightly closer than Badjcinus and the numbat Myrmecobius (baraminic distance 0.34). The five didelphids are definitely separated from the cluster of dasyurids (average baraminic distance 0.44). As with the MDS results for the later Voss and Jansa (2009) character set, the 3D stress for this analysis was poor (0.206), with the minimum stress of at eight dimensions. Discussion Based on the single analysis of procyonids using Baskin s (2004) character matrix, it is difficult to draw any firm conclusions. Baskin s characters are entirely craniodental and not holistic. The BDC results suggest a discontinuity between Amphictis/Broiliana and the remaining procyonids, but the MDS clustering suggests the entire set of procyonids in Baskin s matrix are all part of the same group. In their analysis of the equids, Cavanaugh et al. (2003) found that the ends of a single trajectory of taxa would be negatively correlated when compared directly, even though they were manifestly part of the same linear cluster in the ANOPA results. Given the linear shape of the cluster of procyonid taxa in the MDS results, the negative BDC between Amphictis/Broiliana and the Procyonini is likely the result of being on opposite ends of the linear cluster rather than a genuine discontinuity. We might very tentatively identify the procyonids as a monobaramin, since they appear to form a continuous cluster in MDS, but this will need to be confirmed by additional analyses. In contrast, the three BDC and MDS analyses presented here for the didelphids suggest that Didelphidae may be a holobaramin. The earlier matrix of Voss and Jansa (2003) appears to support a discontinuity between tribe Didelphini and the remaining didelphids, but the bootstrap values for the significant, negative BDC are lower than 90%, indicating that the correlations are only observed with particular combinations of characters. The later Voss and Jansa (2009) character matrix includes non-didelphid outgroups, which has a two-fold effect. First, the didelphids appear as a single, united cluster in both the BDC and MDS results. Second, discontinuity can be inferred between the didelphids and the peramelid/caenolestid outgroup species from the significant, negative BDC with bootstrap values >90%. Likewise, the MDS results for the same baraminic distance matrix show clearly separate clusters of didelphids and nondidelphids. The exceptions are the microbiotheriid Dromiciops and the dasyurids Sminthopsis and Murexia. In the BDC results for the later Voss and Jansa (2009) character matrix, the frequent positive BDC between Dromiciops and the didelphids and its position in the didelphid cluster in the MDS results suggest that Dromiciops belongs to the Didelphidae holobaramin. Indeed, Dromiciops was once part of the didelphid family and only recently has been separated based on anatomical and genetic evidence (e.g., Kirsch et al. 1991; Hershkovitz 1992; Asher et al. 2004). In contrast, the two dasyurids show significant, positive BDC with only 10 of the 44 didelphids, and the MDS results indicate that the dasyurids are separated from the main didelphid cluster. This implies that the dasyurids should not be considered as part of the didelphid cluster. The hypothesis of discontinuity between the didelphids and dasyurids can be evaluated using the character set of Wroe et al. (2000). The five didelphids of that dataset are all negatively correlated with the dasyurids, and the MDS results confirm the separation of these two groups of taxa. The dasyurids themselves are positively correlated and form a single cluster in the MDS results. Thus, we may tentatively identify Dasyuridae as a holobaramin that is discontinuous with the didelphids. Altogether, then, the evidence presented here supports Didelphidae + Microbiotheriidae as a holobaramin. The negative BDC observed between Didelphini and other didelphids with the earlier Voss and Jansa (2003) character set has low bootstrap values and would be superseded by the better supported positive BDC observed among all the didelphids with the later Voss and Jansa (2009) character set. In the later character set, the limited positive BDC with low bootstrap values that seems to support including the dasyurids in the didelphid holobaramin would be contradicted by the clustering pattern in the MDS results and the BDC analysis of dasyurid characters compiled by Wroe et al. (2000). Since both Voss and Jansa (2003, 2009) datasets include whole body characters, they would fit the baraminological ideal of holistic data better than a set of only craniodental characters, as in the case of the procyonids. Still, any sample of characters is not objectively holistic, but these results are broadly consistent with previous analyses of mammal baramins that identified families or subfamilies as baramins (Wood 2008b). The differences between the BDC and MDS analyses of these two taxonomic groups are especially instructive on the interpretation of statistical baraminology results. In early and even recent statistical baraminology studies (e.g., Cavanaugh and Wood 2002; Cavanaugh et al. 2003; Wood 2008b), baramins have been identified based on analyses of a single character matrices, some of which can hardly be called holistic. Similarly, we now know after a decade of studies that positive or negative correlation between taxa can result from factors unrelated to their baraminic status (e.g., Cavanaugh et al. 2003, Wood 2005). These later results call into question the apparently simple interpretation of holobaramins or monobaramins based on a single dataset. JCTS B: Life Sciences Volume 4:17

7 A better approach to identifying baramins using statistical approaches would be the use of multiple datasets representing different samples of taxa and characters. Wood (2005) used an early version of this approach with his multidimensional scaling analysis of three turtle datasets, and Wood (2010) later produced a more comprehensive analysis of the hominin holobaramin based on eight datasets. These types of multi-dataset baraminology studies combined with bootstrapping (Wood 2008a) should become the standard approach for statistical baraminology in the future. In the present study, we should therefore be hesitant in making any firm conclusions about the procyonids since there is only one craniodental dataset with ambiguous BDC and MDS results. With the didelphids, however, we have examined them from two perspectives: that of the didelphids in Voss and Jansa (2003, 2009) and that of the dasyurids in Wroe et al. (2000). These two perspectives lend greater support to the hypothesis that Didelphidae + Microbiotheriidae is a holobaramin. References Asher, R.J., I. Horovitz, and M.R. Sánchez-Villagra First combined cladistic analysis of marsupial mammal interrelationships. Molecular Phylogenetics and Evolution 33: Baskin, J.A Bassariscus and Probassariscus (Mammalia, Carnivora, Procyonidae) from the Early Barstovian (Middle Miocene). Journal of Vertebrate Paleontology 24: Bininda-Emonds, O.R.P Phylogenetic position of the giant panda. In: Lindburg, D. and K. Baragona, eds. Giant Pandas: Biology and Conservation. University of California Press, Berkeley, pp Cavanaugh, D.P. and T.C. Wood A baraminological analysis of the tribe Heliantheae sensu lato (Asteraceae) using Analysis of Patterns (ANOPA). Occasional Papers of the BSG 1:1-11. Cavanaugh, D.P., T.C. Wood, and K.P. Wise Fossil Equidae: a monobaraminic, stratomorphic series. In: Ivey, R.L., ed. 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Jansa Phylogenetic relationships and classification of didelphid marsupials, an extant radiation of new world metatherian mammals. Bulletin of the American Museum of Natural History 322: Wood, T.C A creationist review and preliminary analysis of the history, geology, climate, and biology of the Galápagos Islands. CORE Issues in Creation 1: Wood, T.C. 2008a. Baraminic distances, bootstraps, and BDISTMDS. Occasional Papers of the BSG 12:1-17. Wood, T.C. 2008b. Animal and plant baramins. CORE Issues in Creation 3: Wood, T.C Baraminological analysis places Homo habilis, Homo rudolfensis, and Australopithecus sediba in the human holobaramin. Answers Research Journal 3: Wozencraft, W.C Order Carnivora. In: Wilson, D.E. and D.M. Reeder. Mammal Species of the World. Johns Hopkins University Press, Baltimore, pp Wroe, S., M. Ebach, S. Ahyong, C. De Muizon, and J. Muirhead Cladistic analysis of dasyuromorphian (Marsupialia) phylogeny using cranial and dental characters. Journal of Mammalogy 81: JCTS B: Life Sciences Volume 4:18