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Current knowledge on inquiline oak gall wasp diversity from Mexico (Hymenoptera: Cynipidae)
Abstract
ABSTRACT. The inquiline oak gall wasp fauna from Mexico (Hymenoptera: Cynipidae) has been represented by only two species for years: Synergus dugesi Ashmead, 1899 and Synergus filicornis Cameron, 1883 (= S. furnessana Weld, 1913). Here, the current knowledge on the diversity of inquiline oak gall wasps of the genus Synergus Hartig is summarized and the most recent published advances on the study of their diversity are presented. The species known to date are listed and illustrated together with the information regarding their location and biology. New locations for S. filicornis are presented. The state of current research is commented on and the prospects for future studies about Mexican inquiline oak gall wasps (Cynipidae: Synergini, Ceroptresini) are discussed.
RESUMEN. La fauna mexicana de inquilinos de agallas de cinípidos en Quercus L. (Hymenoptera: Cynipidae) estaba representada, hasta hace pocos años, únicamente por dos especies: Synergus dugesi Ashmead, 1899 y Synergus filicornis Cameron, 1883 (= S. furnessana Weld, 1913). En este trabajo, se resume el estado actual del conocimiento de los inquilinos del género Synergus Hartig de agallas de cinípidos en encinos de México y se presentan los avances más recientes sobre el estudio de su diversidad. Se enlistan e ilustran las especies conocidas junto con la información referente a su localización y biología. Se presentan nuevas localidades para S. filicornis. Se comenta el estado de las investigaciones que se están llevando a cabo en la actualidad y se discuten las perspectivas de estudios futuros de los inquilinos mexicanos en encinos (Cynipidae: Synergini, Ceroptresini).
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The inquiline oak gall wasp fauna emerged from agamic galls of
Andricus quercuslaurinus Melika and Pujade-Villar, 2009, a dangerous pest for
Quercus affinis Scheidw. and Q. laurina Humb. et Bonpl. forests in Mexico, was
studied for the first time. Three species in the genus Synergus Hartig, 1840
(Synergini) were obtained from these galls: Synergus davisi (Beutenmüller, 1907),
previously known only from the United States of America; Synergus estradae
Pujade-Villar and Lobato-Vila, 2016, described from the state of Morelos (Mexico);
and Synergus pseudofilicornis Lobato-Vila and Pujade-Villar n. sp., so far
determined as Synergus filicornis Cameron, 1883 (= Synergus furnessana Weld,
1913) and here considered as a new species. Synergus pseudofilicornis n. sp. is
described and illustrated. A redescription and photographs of S. davisi, as well as
new distributions and host associations of S. davisi and S. estradae, are given. The
value of the inquilines as biological control agents for A. quercuslaurinus is
discussed, and a preliminary list of parasitoids obtained from the galls is presented.
Here we describe five new species of Synergus Hartig with partially smooth mesopleurae that were obtained from oak cynipid galls on different species of Quercus L. in Mexico: Synergus cibriani sp. nov., Synergus gilletti sp. nov., Synergus longimalaris sp. nov., Synergus longiscapus sp. nov. and Synergus striatifrons sp. nov. To date, this trait has been observed in at least 18 American species, 16 recorded in North America (United States of America) and only 2 in Central America (Panama): Synergus elegans Nieves-Aldrey and Medianero and Synergus laticephalus Nieves-Aldrey and Medianero. The type material and additional material of Synergus punctatus Gillette from the United States have been examined and compared with S. gilletti sp. nov. The type material and additional material of S. elegans from Panama and Synergus citriformis (Ashmead) from the United States have also been examined and compared. This study records Synergus elegans in Mexico for the first time and classifies it as a syn. nov. of S. citriformis. Descriptions and diagnoses for the new species and a key to their identification and differentiation from their closest related species are given. Their distribution and host associations are also commented on.
Two new species of Synergus Hartig obtained from inconspicuous oak galls inside Quercus rugosa Née acorns (Synergus equihuai Pujade-Villar & Lobato-Vila n. sp.) and from tumor galls collected in Q. obtusata Bonpl. (Synergus estradae Pujade-Villar & Lobato-Vila n. sp.) are described from Mexico. The most important morphological characters are illustrated; in addition, the new species are morphologically differentiated from the closest related species.
Fifteen new species of cynipid inquilines, Saphonecrus chinensis Tang & Schweger, S. gilvus Melika & Schweger, S. globosus Schweger & Tang, S. leleyi Melika & Schweger, S. lithocarpii Schweger & Melika, S. longinuxi Schweger & Melika, S. morii Schweger & Tang, S. nantoui Tang, Schweger & Melika, S. nichollsi Schweger & Melika, S. pachylomai-Schweger, Tang & Melika, S. robustus Schweger & Melika, S. saliciniai Melika, Tang & Schweger, S. shanzhukui Melika & Tang, S. symbioticus Melika & Schweger, and S. taitungi Schweger, Tang & Melika, from the Eastern Palaearctic are described. Descriptions, diagnoses, biology, and host associations for the new species, and a key to Palaearctic Saphonecrus species are given. All new taxa form distinct units as demonstrated by the molecular phylogenetic analyses of Palaearctic Saphonecrus species. The status of some earlier described Saphonecrus species is discussed also. The Synergini genus Lithonecrus Nieves-Aldrey & Butterill, 2014 is synonymized with Lithosaphonecrus Tang, Melika & Bozso, 2013. Three Saphonecrus species are transferred to Synergus: Synergus brevis (Weld) comb. nova, Synergus hupingshanensis (Liu, Yang & Zhu) comb. nova, and Synergus yukawai (Wachi, Ide & Abe) comb. nova. Synophrus vietnamensis Abe, Ide, Konishi & Ueno is transferred to Lithosaphonecrus: Lithosaphonecrus vietnamensis Abe, Ide, Konishi & Ueno), comb. nova. The current number of valid Saphonecrus species worldwide is 36.
Gall wasps (Cynipidae) represent the most spectacular radiation of gall-inducing insects. In addition to true gall formers, gall wasps also include phytophagous inquilines, which live inside the galls induced by gall wasps or other insects. Here we present the first comprehensive molecular and total-evidence analyses of higher-level gall wasp relationships. We studied more than 100 taxa representing a rich selection of outgroups and the majority of described cynipid genera outside the diverse oak gall wasps (Cynipini), which were more sparsely sampled. About 5 kb of nucleotide data from one mitochondrial (COI) and four nuclear (28S, LWRh, EF1alpha F1, and EF1alpha F2) markers were analyzed separately and in combination with morphological and life-history data. According to previous morphology-based studies, gall wasps evolved in the Northern Hemisphere and were initially herb gallers. Inquilines originated once from gall inducers that lost the ability to initiate galls. Our results, albeit not conclusive, suggest a different scenario. The first gall wasps were more likely associated with woody host plants, and there must have been multiple origins of gall inducers, inquilines or both. One possibility is that gall inducers arose independently from inquilines in several lineages. Except for these surprising results, our analyses are largely consistent with previous studies. They confirm that gall wasps are conservative in their host-plant preferences, and that herb-galling lineages have radiated repeatedly onto the same set of unrelated host plants. We propose a revised classification of the family into twelve tribes, which are strongly supported as monophyletic across independent datasets. Four are new: Aulacideini, Phanacidini, Diastrophini and Ceroptresini. We present a key to the tribes and discuss their morphological and biological diversity. Until the relationships among the tribes are resolved, the origin and early evolution of gall wasps will remain elusive.
A new term, agastoparasitism, is proposed for parasitism among closely related species. Cynipid inquilines are typical agastoparasites. They cannot induce galls; instead their larvae live inside the galls formed by other cynipids. As in many other groups of agastoparasites, there are two competing hypotheses for the evolutionary origin of cynipid inquilines: either they arose from one of their cynipid hosts, and later radiated to exploit other gall-inducing cynipids (monophyletic origin), or they arose repeatedly, each inquiline from its host (polyphyletic origin). These hypotheses for the origin of cynipid inquilines were tested by a phylogenetic analysis of representative species of cynipid gall inducers and inquilines based on adult morphological characters. The analysis supported the monophyly of the inquilines and indicated an origin from gall inducers related to the genus Diastrophus, one of the current host groups. To examine whether the result of the analysis was influenced by convergent similarities among inquilines because of their similar mode of life, all putative apomorphies shared by some or all of the inquilines but not occurring in any of the gall inducers were removed. Despite this, the phylogenetic conclusions essentially remained the same, that is, the support for inquiline monophyly was not caused by convergent evolution. Based on these results, adaptive aspects of the evolutionary origin and maintenance of cynipid inquilinism are discussed, as well as general patterns in the evolution of agastoparasitism.