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Almost but not quite a subspecies: a case of genetic but not morphological diagnosability in Nicrophorus (Coleoptera: Silphidae)

Biological journal of the Linnean Society 2011 v.102 no.2 pp. 311-333
DNA barcoding, Nicrophorus, allopatry, discriminant analysis, genes, genetic distance, genetic variation, mitochondrial DNA, morphometry, new species, nucleotide sequences, phylogeny
Allopatric populations that show genetic differentiation but lack phenotypic diagnosability are difficult to classify. In 1946, Arnett described a new species of burying beetle (Silphidae: Nicrophorus) from Luzon Island, Philippines, Nicrophorus benguetensis. In 2002, Sikes et al., finding the species to be inconsistently diagnosable, synonymized N. benguetensis under Nicrophorus nepalensis Hope 1831. We rigorously and quantitatively test the validity of N. benguetensis using several different species delimitation criteria. We employed discrete and quantitative character-based methods to test similarity criteria using linear morphometrics in a discriminant analysis framework, and percentage sequence divergences based on sequences from four mitochondrial DNA (mtDNA) genes (COI, COII, ND4 and ND5) and two nuclear genes [28S (D2 region) and CAD]. We also employed tree-based methods to test phylogenetic criteria using mtDNA sequences and morphology with parsimony, Bayesian inference and maximum likelihood. The analysis of discrete phenotypic characters did not reliably diagnose N. benguetensis. The results of the discriminant analysis provided moderate support for the validity of N. benguetensis (71.4% of the specimens were properly classified). There is adequate genetic distance between N. benguetensis and its nearest neighbour for DNA barcoding to identify an unknown sequence, although this may be an artefact of the small sample size. Phylogenetic analyses of the morphological data, with and without the morphometric data, yielded unresolved trees. Molecular phylogenetic results found N. benguetensis to be monophyletic, but neither clearly rejected nor supported its validity. Two of our approaches using molecular data diagnosed N. benguetensis, but those that used phenotypic data did not exceed a 75% success rate. We conclude, therefore, that N. benguetensis, despite being a weakly distinct allopatric population, and deserving of additional study, should remain synonymized until further analysis suggests otherwise.