CO1 and ITS alignments of bladder grasshopper species

Polymorphism in one sex often occurs amongst arthropods, and is often
linked with alternate mating strategies. Bladder grasshoppers show
distinct polymorphism resulting in two different male morphs utilizing two
different mating strategies. Primary males are larger in size and make use
of acoustic communication to attract mates, while secondary (alternate)
males are significantly smaller in size, flightless, and do not call,
making use of a “sneaker” or satellite strategy. Three species of bladder
grasshoppers have been described (Parabullacris vansoni, Paraphysemacris
spinosus, and Pneumoracris browni) that only have an alternate male morph
and no primary male, whereas all other species in the family are described
as having only a primary male. However, it is now known that both types of
males occur in at least three species (Bullacris discolor, B.
membracioides, and B. obliqua). Thus, the validity of the taxonomic
descriptions for species with no identified primary male has come into
question. We therefore conducted a phylogenetic analysis to examine the
genetic relationships between primary and secondary males for all species
where secondary males are documented or suspected to occur. This was done
using both mitochondrial (COI) and nuclear (ITS) DNA markers. We confirm
that dimorphic males are indeed conspecific for the species observed to
have both primary and alternate males. We also demonstrate that
Parabullacris vansoni, Pneumoracris browni and Paraphysemacris spinosus
are invalid taxa as they are conspecific alternate male morphs of B.
unicolor, Peringueyacris namaqua and Physemacris variolosa, respectively.

节肢动物中常存在单一性别的多态现象（polymorphism），且该现象往往与交替交配策略相关联。泡蜢（bladder grasshoppers）呈现出显著的多态现象，产生两种不同的雄性形态，各自采用截然不同的交配策略。初级雄性体型更大，通过声学通讯吸引配偶；而次级（替代）雄性体型显著更小、无飞行能力且不发出鸣唱，采用“偷袭者”（sneaker）或“卫星”交配策略。目前已正式描述的3种泡蜢（Parabullacris vansoni、Paraphysemacris spinosus及Pneumoracris browni）仅存在替代雄性形态，无初级雄性；而该科其余所有物种均被认为仅具有初级雄性。但目前已有研究证实，至少有3个物种（Bullacris discolor、B. membracioides及B. obliqua）同时存在两种雄性形态。因此，那些未被发现存在初级雄性的物种的分类学描述有效性受到了质疑。为此，本研究针对所有已记录或疑似存在次级雄性的物种，开展了系统发育分析，以探究其初级与次级雄性间的遗传亲缘关系。本研究采用线粒体（COI）与核（ITS）DNA标记开展分析。本研究证实，在同时存在两种雄性形态的物种中，二态雄性确实为同一物种。本研究同时证实，Parabullacris vansoni、Pneumoracris browni及Paraphysemacris spinosus均为无效分类单元，它们分别是B. unicolor、Peringueyacris namaqua及Physemacris variolosa的次级雄性形态。