For malaria parasites to be transmitted from human to human requires at least two mosquito bites. First, to become infected the mosquito needs to feed on an infectious human host. The first bloodmeal that can potentially infect a mosquito is usually consumed 2 days after mosquito emergence from its aquatic breeding site 
, but this is likely to be temperature-dependent 
. To then onwardly transmit, the mosquito needs to take another bloodmeal when it is infectious. Transmission to a new host cannot occur until the Extrinsic Incubation Period (EIP) of the parasite is completed. The EIP is the length of time it takes the malaria parasite to complete its development within the mosquito and migrate to the salivary glands, and is one of the key rate limiting steps in the transmission of malaria. EIP is known to be strongly temperature sensitive, taking anywhere from around 10 days to >30 days depending on conditions 
Human malaria mosquitoes (Anopheles
spp.) generally exhibit discrete feeding cycles in which blood feeding only occurs at the beginning of a gonotrophic cycle and then not again until the blood has been digested, a batch of eggs has matured and oviposition is completed (e.g. 
). While there is the possibility of a small proportion of Anopheles
mosquitoes taking multiple blood feeds within a cycle 
, evidence suggests this behavior might be restricted to smaller (and more energy depleted) mosquitoes during the first feeding cycle, immediately after emergence from their breeding site 
In most cases blood feeding is coupled to reproduction, and therefore the frequency of possible transmission events (i.e. acquiring the parasite in an early feeding event and then passing it on in a later feed) depends on the duration of this gonotrophic cycle. Again, the length of the gonotrophic cycle is strongly temperature dependent 
. Under warmer conditions (~30°C), the gonotrophic cycle can be completed in just 2–3 days 
resulting in a high frequency of blood feeding. Under cooler conditions (15–20°C), on the other hand, blood feeding might occur only once every 6–13 days 
While both parasite development and the gonotrophic cycle are strongly affected by temperature, these processes are independent of one another and the nature of their temperature dependence differs (see results
). There exists the possibility, therefore, that the EIP and feeding could be out of phase such that the parasite could complete development at an early or mid-point of a gonotrophic cycle. In this case, the parasite has a ‘waiting period’ where it cannot be transmitted until the next feed, even though the time since the initial infected bloodmeal has extended beyond the EIP.
Widely used malaria transmission models, such as the Ross-MacDonald models (see 
for an overview of this family of models and 
for illustrative applications), do not consider delays in infection due to a pre-bloodmeal period, or possible delays in onward transmission due to asynchrony between parasite development and the gonotrophic cycle. A limited number of models have included such delays, but have not explored the implications for transmission explicitly 
. The aim of the current study, therefore, is to address this knowledge gap.
We begin with some illustrative, proof-of-principle empirical investigations using the Asian malaria vector, Anopheles stephensi, and a rodent malaria, Plasmodium yoelii, to explore the assumptions that the pre-blood meal period is temperature-dependent, blood feeding is linked to the gonotrophic cycle, and that the EIP and the gonotrophic cycle are affected differentially by temperature. We then extend the study using some simple modifications of an established model for vectorial capacity. Using previously defined relationships between temperature and the gonotrophic cycle, and temperature and the EIP of the human malaria P. falciparum, we explore how biting and EIP can move in and out of phase across different thermal environments. This effect, combined with an initial temperature-dependent pre-bloodmeal period, can lead to substantial reductions in transmission intensity.